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CANINE FUNCTION IN SM1LODON
(MAMMALIA; FELIDAE; MACHAIRODONTINAE)

William A. Akersten

Natural History Museum of Los Angeles County • 900 Exposition Boulevard « Los Angeles, California 90007

SERIAL PUBLICATIONS OF THE
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The scientific publications of the Natural H istory Museum of Los Angeles County have been
issued at irregular intervals in three major series; the articles in each series are numbered
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® Contributions in Science, a miscellaneous series of technical papers describing orig-
inal research in the life and earth sciences.

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in the life and earth sciences. This series was discontinued in 1978 with the issue of
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in Science.

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SCIENTIFIC PUBLICATIONS COMMITTEE

Craig C. Black, Museum Director
Donald C'haput

Daniel M. Cohen, Committee Chairman
John M. Harris
Charles L. Hogue

Robin A. Simpson. Managing Editor
Gary D. Wallace
Edward C. Wilson

CANINE FUNCTION IN SMILODON
(MAMMALIA; FELIDAE; MACHAIRODONTINAE)

William A. Akersten'

ABSTRACT. A number of sabertoothed mammal features, exem-
plified by Smilodon from Rancho La Brea, cannot be reconciled with
hypotheses that the upper canines were used for stabbing or slashing:
the relatively dull sabers which would require enormous force to
penetrate the hide of a prey animal, the robust mandible with func-
tional but small canines, the geometric relationships of the skull and
the reconstructed head-depressing musculature, and the presence of
canine opposition (necessary for biting but not for stabbing or slash-
ing). It is also difficult to envision intermediate steps of behavior
and morphology in the multiple evolution of stabbing or slashing
from biting ancestors. These apparent anomalies can be resolved
(and other features can be explained) by hypothesizing that the upper
canines were employed in a shearing, as opposed to puncturing, bite
accomplished by depressing the cranium against immobilized man-
dibles. The probable area of attack was the abdomen. Most, if not
all, other sabertoothed mammals appear to possess canine opposi-
tion and probably employed some variation of the attack method-
ology suggested for Smilodon. Several lines of evidence suggest that
Smilodon possessed some degree of cooperative social behavior. The
attack methods of the extant Komodo dragon, Varanus komodoen-
sis, appear to be analogous to those hypothesized for Smilodon and
the extinct giant varamds of Australia may have preempted the niche
occupied by sabertoothed mammals on other continents.

INTRODUCTION

The suite of unusual specializations which typify saber-
toothed mammals has intrigued researchers for decades.
Practically every possible mode of saber use has been pos-
tulated, including using them as “can-openers” for glypto-
donts (Brandes, 1900) or for slicing carrion (Bohlin, 1940).
Matthew (1901) refers to earlier suggestions (without cita-
tion) that sabertooths used their canines to grub for marine
molluscs as walruses supposedly do, as tree climbing aids,
or (in forms with reduced mandibular flanges) to stab with
the mouth closed.

More recent studies (e.g.. Hough, 1949; Miller, 1969; Gon-
yea, 1976; Martin, 1980; and Emerson and Radinsky, 1980)
have firmly established sabertooths as active predators well
adapted to catching prey and eating meat. Presently accepted
hypotheses concerning saber function usually center about
their use in stabbing and/or slashing, probably at the throat

Contributions in Science, Number 356, pp. 1-22
Natural History Museum of Los Angeles County, 1985

of their prey. While these interpretations appear to be con-
sistent with many sabertooth specializations, other consid-
erations have led me to question the efficacy of this type of
attack.

One difficulty I have with accepting a stabbing or slashing
attack by Smilodon involves the problem of forcing a pair
of such weapons into the hide and flesh of a prey animal. As
compressed and seemingly sharp as the sabers may be, they
are far from being equivalent to a steel knife. Forcing even
a slightly dulled steel knife through the hide of a large mam-
mal can be very difficult. Other mammalian carnivores with
conical canines use the opposing force of the lower canines
to facilitate penetration of the upper canines (and vice versa)
during an attack. If they simply tried to force the upper
canines into their prey without using the lower canines, they
would fail. Similarly, if Smilodon were to try to stab or slash
without an opposing force, considerable momentum would
have to be built up by the time the sabers contacted the prey.
I believe that the problem of developing enough force to
drive a pair of rather dull sabers into hide and flesh raises
serious questions about any stabbing or slashing hypothesis.

An important theoretical point involves the fact that sa-
bertooth adaptations arose independently within at least four
different groups of mammals— borhyaenid marsupials, creo-
donts, nimravids, and felids. As the normal kill method in
all living mammalian carnivores (and, presumably, all ances-
tors of sabertoothed types) is the canine bite, postulating a
slash or stab requires the multiple, convergent development
of a radically different method of kill behavior. I find it
impossible to envision the necessary intermediate combi-
nations of morphology and behavior that would be re-
quired by a gradualistic model of evaluation and equally
impossible to accept that such a complex combination of
changes could occur even once within the framework of punc-
tuated models.

These and other difficulties with the slashing/stabbing hy-

1. Section of Vertebrate Paleontology, Natural History Museum
of Los Angeles County, Los Angeles, California 90007.

ISSN 0459-8113

potheses raised the question of whether the peculiar spe-
cializations of sabertoothed mammals have resulted in more
radical interpretations of saber function than is necessary. In
an effort to address this question, I studied the large Rancho
La Brea sample of Smilodon in the collections of the George
C. Page Museum (specimen prefixes LACMHC and
LACMRLP), a branch of the Natural History Museum of
Los Angeles County (LACM).

MORPHOLOGY

Three character complexes were most important in the de-
velopment of the various slashing/stabbing hypotheses: (1)
the apparently weak mandible with a reduced canine which
does not appear to have been of much use during an attack,
(2) the evidence for powerful head-depressing muscles which
could be used in a stabbing or slashing action, and (3) the
striking hypertrophy of the upper canines which immediately
conjures up images of the edged weapons man uses to slash
or stab. Re-examination of these characters and consider-
ation of other morphologic features yield additional infor-
mation not always in agreement with previous interpreta-
tions.

OVERALL STRUCTURE

The Smilodon from Rancho La Brea approximated the size
of a modem African lion but had very powerful forelimbs,
a proportionately larger head, and a short tail. The forelimbs,
with their large, retractile claws (Gonyea, 1976), would have
been very well suited to pulling down and immobilizing fairly
sizable prey animals. Limb and foot structure and propor-
tions (Merriam and Stock, 1932; Gonyea, 1976; Martin, 1980;
Shaw and Tejada-Flores, in press) and vertebral structure
(Merriam and Stock, 1932; Slijper, 1946) indicate that Smi-
lodon was not cursorial but probably was capable of short,
rushing attacks. Most recent authors generally agree that
Smilodon probably stalked close to its potential prey, then
rushed a short distance from ambush. I believe that the mas-
sive forelimbs were employed in grasping the upper part of
the prey’s body from the side and pulling it down towards
the attacker so that the abdomen would be exposed on the
opposite side. This method of bringing down prey is similar
to that often used by many of the larger living felids when

attacking large prey except that they may also employ their
teeth in the attack (Leyhausen, 1979; Schaller, 1967, 1972).

MANDIBULAR COMPLEX

The greatly reduced coronoid process has been repeatedly
used as evidence for a relatively weak bite (e.g., Matthew,
1910; Merriam and Stock, 1932) because it would have pro-
vided less leverage for the temporalis muscle. Emerson and
Radinsky (1980), elaborating on the work of Kurten (1952),
showed that the probable bite strength during camassial oc-
clusion was roughly equivalent to that of comparably sized
living felids, but at full gape, it was undoubtedly rather weak.
This resulted not only from the poor lever arm for the tem-
poralis but also from the reduction of the masseter. They
also determined that the relative cross-sectional area of the
mandible below the camassial was about equivalent in sa-
bertooths and living felids. The diastemal region superficially
appears weaker in Smilodon because it is relatively shallow.
However, Figures 1 A, B compare cross sections cut through
the shallowest part of the diastema in a young adult African
lion (Panthera leo) and a young adult Smilodon of about the
same size. Not only is the relatively greater breadth of this
area in Smilodon readily apparent, but the outer layer of
compact bone is far thicker. If anything, the mandible of
Smilodon appears much stronger than that of the lion. Sev-
eral longitudinally sectioned and many broken mandibles in
the Rancho La Brea collections at the George C. Page Mu-
seum demonstrate that this very strong internal construction
characterizes the entire Smilodon mandible. In overall ap-
pearance, the Smilodon mandible is stouter than that of the
lion with a massive, very rugose symphyseal region (Fig. 1C)
most similar to that of Panthera among the type III sym-
physes described by Scapino (1981). The major differences
between the two appear in the outline of the symphyses and
the distribution of the largest rugosities. In Smilodon these
occur toward the ventral portion at or near the ventral tu-
bercle while in Panthera they are found near the posterior
margin. The evidence strongly indicates that powerful forces
impinge upon the mandible and that it was not simply used
for shearing at the camassials or pulling flesh off prey in
conjunction with the incisors, the only other postulated uses
for the mandibular dentition (Emerson and Radinsky, 1980;
Miller, 1969).

Forces acting on the mandible have major components
transmitted through the condyloid process to the glenoid

Figure 1. Panthera leo. A. Cross section through diastema of mandible, LACMRLB JGT2. Rest are Smilodon. B. Cross section through
diastema of mandible, LACMHC 7108. C. Stereopair of left mandibular symphysis, LACMRLP R1 1258. D. Ventral view of anterior mandible,
LACMHC 2001-2, showing anterior projection of flanges. E. Stereopair of right mastoid process, LACMHC 2001-2 (retroarticular process at
bottom). F, G, H. Posteroventral views of left portion of cranium, LACMRLP 20273, and associated atlas, LACMRLP 20276, depicting
relationships of atlas and mastoid process. F. Cranium only. H. Cranium with atlas articulated and rotated to extreme anteroventrad position.
G. Same as H but with “ghost image” of ammonium chloride coated atlas in double exposure. Note the alignment of portions of the lateral
margins of atlas wing and mastoid process between white marks in G. Scales: A, B, F, G, H, scale bar = 2 cm (bar between A and B applies
to both and bar on H also applies to F and G). C, D, E in cm.

2 Contributions in Science, Number 356

Akersten: Canine Function in Smilodon

Contributions in Science, Number 356

Akersten: Canine Function in Smilodon 3

fossa of the squamosal. This region independently reflects
the gross magnitude of such forces. In comparison to the
African lion, the neck of Smi/odon's condyloid process is
much stouter and the condyle is wider with a considerably
larger articular surface. In the cranium of Smilodon, the ret-
roarticular process is thick and stout, the entire zygoma is
very well developed and the base of the zygomatic process
of the temporal provides massive support for the glenoid
fossa. The entire articular complex and supporting structures
demonstrate again that the mandible of Smilodon was sub-
jected to very strong forces during some portion of its use.

While the lower canines of Smilodon are relatively small
in comparison to body size, they are far from vestigial. They
approximate the size of canine found in a medium-sized
mountain lion (Felis concolor) but they appear even smaller
because of their proximity to the very large incisors. The
lower canines are stout, sharp, and recurved posteriorly with
strong roots and relatively thick enamel. Their posterior and
medial margins bear sharp, finely serrate ridges (see Merriam
and Stock, 1932: pi. 13, figs. 8, 8a). While the medial serrate
ridge may have functioned with the incisor battery in wor-
rying loose chunks of meat as described by Miller (1969),
the only apparent function for the posterior serrate ridge
would have been to facilitate puncturing. Thus, the reduced
lower canines of Smilodon were still capable of functioning
like those of living felids in penetrating the hide and flesh of
prey animals, but would not have produced as large or deep
wounds as the lower canines of an equivalent-sized true cat.
The large roots and the heavy bone surrounding them are
evidence that the canines were also subjected to considerable
stress.

One often discussed feature of the mandible is the antero-
ventral mandibular flange. The ventral development of this
flange in Smilodon is relatively slight, when compared to
many other sabertoothed mammals, and quite variable. It
has been suggested that a highly developed ventral flange
served to protect the sabers from breakage or to protect the
neck from accidental injury by the sabers (Scott and Jepsen,
1936; Martin, 1980). A more plausible explanation has re-
cently been advanced by Dawn A. Adams and Daniel B.
Adams (pers. comm., 1983), which relates the ventrad de-
velopment of the flange to a powerful digastric musculature
required by forms with nearly vertical occiputs. Several stud-
ies (e.g., Schultz et ah, 1970) have suggested that the flange
became reduced in Smilodon to allow stabbing with the mouth
closed. This appears to be improbable because there would
then be no function for that portion of the posterior serrations
on the upper canine which extend dorsad to the ventral bor-
der of the closed mandible all the way to the gum line. Fur-
thermore, if the sabers were used to stab with the mouth
closed, the proximal serrations should wear smooth more
slowly than the distal serrations. None of the La Brea spec-
imens display such differential wear. Also, a major potential
problem would result from the shock transmitted to occlud-
ing upper and lower teeth (particularly the interlocking in-
cisors) when the mandible struck the prey in a closed mouth
stab.

In contrast to the moderate and variable ventral devel-

opment of the flanges, they are consistently well developed
anteriorly. The anterior portions of the flanges flare laterally
and become slightly thickened at their margins. In ventral
view, the anterior outline of the symphyseal region is strongly
concave with the symphysis situated at the axis of a consid-
erable depression (Fig. ID). As described below, I believe
that the anterior development of the flange played an im-
portant part during Smilodon' s attack.

INCISOR BATTERY

The basic form and function of the upper and lower incisors
have been well described by Merriam and Stock (1932) and
Miller (1969). Overall, they are very large, sharp, recurved
posteriorly, and bear huge roots. The bone anterior and pos-
terior to the alveoli is very stout. The incisor battery appears
very prognathous with the sharp apices of the upper and
lower incisors (and the lower canines) completely interlock-
ing when the jaw is closed (Fig. 2). All tend to bear ridges
on their medial and lateral margins. These ridges are usually
very finely serrate in unworn teeth with the serrations be-
coming progressively better developed from the medial (where
they may be absent) to the lateral incisors. The ridges curve
posteriad toward the bases of the crowns to form cingula
which frequently bear small cuspules. Placement of the cin-
gular cusps is such that they provide additional shear against
the tips of the opposing incisors when the jaw is completely
closed. If the incisors were used to gnaw flesh from bones as
suggested by Miller (1969), one would expect to find wear
on the anterodistal portions of the crowns. None of the ex-
amined specimens exhibited such wear. As noted by Merriam
and Stock (1932) and further elaborated by Miller (1969),
the incisor battery, including the lower canines, forms an
immensely powerful puncturing and gripping device. I would
add that it was also capable of developing considerable shear
between opposing teeth.

HEAD-DEPRESSING MUSCULATURE

A number of osteological characters indicative of powerful
head-depressing muscles in various sabertoothed mammals
have been cited as evidence supporting a stabbing or slashing
attack (Matthew, 1910; Merriam and Stock, 1932). Emerson
and Radinsky (1980) question these interpretations, noting
that the insertion scars for the rectus capitus ventralis and
longus capitus (at the basioccipital-basisphenoid suture) are
not unusually well developed and asserting that “the ster-
nomastoid and the cleidomastoid do not leave discrete scars
on the mastoid process.” While it is true that the insertion
scars for the rectus capitus ventralis and the longus capitus
are only modestly developed in Smilodon, the broad tip of
the mastoid process is very rugose with several deep pits (Fig.

1 E) which may coalesce to form a large groove. I agree with
Merriam and Stock (1932:33) that this area probably served
as the insertion for well-developed stemomastoid and clei-
domastoid muscles. In addition, the manubrium of the ster-
num (origin of the stemomastoid) is relatively robust in Smi-
lodon. The enlarged mastoid processes would give these
muscles considerably more leverage to depress the head than

4 Contributions in Science, Number 356

Akersten: Canine Function in Smilodon

in typical felines. Another possible explanation for at least a
portion of the apparent muscle scars at the tip of the mastoid
process is that they served as the area of origin for a very
well developed digastric musculature (Dawn A. Adams and
Daniel B. Adams, pers. comm., 1983).

There is evidence for additional, and even more powerful,
head-depressing muscles in Smilodon. The entire postero-
lateral portions of the enlarged mastoid processes appear to
be covered with very large muscle scars (Fig. IE). Matthew
(1910) believed that this area provided attachment for the
sternomastoid which had shifted from its normal attachment
area at the tip of the mastoid process. Flowever, if the atlas
of Smilodon is rotated anteroventrally about the atlantooc-
cipital articulation, the parts of the lateral margins of its wings
not only align quite well with posterolateral muscle scars of
the mastoid processes but would also effectively block other
muscles of posterior origin which might potentially insert on
these scars (Figs. 1 F, G, FI). Modem felids and canids possess
tiny mastoid processes with only very minute muscles ex-
tending between them and the atlas wings. Descriptions of
modern felid and canid anatomy often ignore, or are not
consistent in the terminology and interpretations of, these
muscles. Though quite different in detail, the highly devel-
oped mastoid processes of the giant panda, Ailuropoda, more
closely approximate those of Smilodon and their anatomical
relationships have been thoroughly described by Davis (1964).
He states that the rectus capitus lateralis inserts on the “pos-
terior surface of the mastoid process near its outer edge” (p.

1 70). The obliquus capitus anterior is described as inserting
“just above the mastoid process” (p. 170) but on his figure
20, its insertion is shown as including much of the posterior
surface of the mastoid process. Both muscles originate on the
ventral surface of the tip of the atlas wing which, in Smilodon,
is posteriorly elongated and deflected ventrad (Figs. 3A, B;
5A). Thus, it appears highly probable that very well devel-
oped muscles (which may or may not be homologous with
those of Ailuropoda) extended from the ventral surfaces of
the atlas wings to the posterolateral margins of the mastoid
processes. The enlargement of the atlas wings and mastoid
processes in Smilodon would give this atlantomastoid mus-
culature increased length and considerable leverage to de-
press the head about the atlantooccipital articulation. I con-
sider them to have been more important in this action than
other head-depressing muscles. Additional evidence for the
presence of powerful muscles originating from the ventral
portion of the atlas wings is provided by the large attachment
area on the axis for the obliquus capitus posterior, which
originates from the entire neural arch and inserts on the
dorsal surface of the atlas wings (Davis, 1964). This muscle
functions to rotate the atlantoaxial articulation and prevents
the atlas wings from being drawn anteriad during contraction
of the atlantomastoid musculature. In comparison to Pan-
thera, the neural arch of the axis in Smilodon (Fig. 3C) is
proportionately larger and extends more posteriad relative
to the centrum indicating a well-developed obliquus capitus
posterior. Reconstructions of the major head-depressing
muscles are depicted in Figure 5A. Overall ventrad move-
ment of the head and neck would have been aided by a

powerful scalene musculature indicated by the enlarged
transverse processes on the cervical vertebrae (Matthew,
1910).

All available evidence leads to the almost inescapable con-
clusion that Smilodon possessed strongly developed muscles
for head depression, especially those which rotate the atlan-
tooccipital articulation. While these have been used as pow-
erful evidence for a stabbing or slashing mode of saber use,
it is not impossible that powerful head depression could have
served some other purpose, as will be proposed below.

UPPER CANINES

The saber morphology of Smilodon has been well described
by Merriam and Stock (1932). To review briefly, they are
long, slender, recurved, blade-like teeth with an extremely
thin veneer of enamel (Fig. 4). The enamel extends to the
gum line along the posterior margin but its extent is variable
on the rest of the tooth, averaging only about two thirds as
far. Very fine enamel serrations extend from the tip to the
gum line on the posterior margin and, variably, between one
half to two thirds as far from the tip on the anterior margin.
The posterior margin is more compressed than the anterior,
especially toward the base of the exposed portion. Here, the
anterior margin becomes quite rounded proximal to the ter-
mination of the anterior serrations. The serrations are fre-
quently worn, even smooth, in older individuals. Although
this wear indicates that the canines were not used simply for
display, 1 believe that the fineness of the serrations and the
very thin enamel layer covering the saber in Smilodon dem-
onstrate relatively infrequent use (such as during the kill but
not regularly during feeding). More frequent use, resulting in
more severe wear, should have required thicker enamel and
strongly developed serrations. The large root of the upper
canine, about 40% of its total length, undoubtedly reflects
the need for support during strenuous use.

The bone surrounding the canine alveolus is relatively thin
along the medial surface, but this area is braced to some
extent by the palatal portions of the maxillary and premax-
illary bones. On the lateral surface, it is thin at the alveolar
margin but thickens a little toward the root region, especially
adjacent to the long axis of the alveolus. Heavier bone occurs
along the anterior and posterior margins of the alveolus,
indicating more anterior and posterior stress on the canine.
In individuals with fully developed canines, very thick bone
surrounds the pointed tip of the canine root and extends
from the tip along the anterior margin for a short distance.
The canine root does not extend as far posteriad as the convex
surface on the external portion of the maxillary suggests, but
terminates above or slightly anterior to the infraorbital fo-
ramen. Younger specimens, in which the canine root is still
forming, have alveoli with broadly U-shaped terminations
surrounded by relatively thin bone. These canine alveoli also
extend farther posteriad, almost to the posterior margin of
the maxillary bone. Thus, thickening of the bone is accom-
plished by a filling in of the alveolus as the root becomes
fully formed. The mass of bone at the tip and along the
proximal anterior edge of the root is consistent with the need

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Akersten: Canine Function in Smilodon 5

Figure 2. Anterior view of Smilodon skull and mandible, LACMHC
2001-1, showing interlocking of incisors. Scale bar = 3 cm. From
Merriam and Stock (1932: pi. 2, fig. 3) with permission, Carnegie
Institution of Washington.

to provide support for fairly strong forces developed by push-
ing the tip against a resistant object. If the sabers were used
regularly in anterior or posterior slicing actions, I would ex-
pect to find more bone supporting the distal portions of the
anterior and posterior alveolar margins.

The importance of the sabers to Smilodon is illustrated by
the similar morphology of the deciduous upper canines which
are, however, concave on their medial surfaces in order to
accommodate considerable eruption of the permanent ca-
nines before the deciduous canines are lost (Merriam and
Stock, 1932). Unlike modem felines, in which there is a brief
period of time when the deciduous canines have become too
weak to use and the permanent canines have not yet erupted

to the point of becoming functional (Leyhausen, 1979), Smi-
lodon always maintained functional upper canines (Tejada-
Flores and Shaw, 1984).

As briefly, mentioned in the introduction, the supposed
knife-like appearance of the sabers is only valid when com-
pared to a typical conical carnivore canine, not when com-
pared to a genuine steel knife. Cross sections through a typical
saber (Fig. 4) more nearly resemble somewhat flattened ovals
than sections through a metal blade. The very tips of the
upper canines, which would initiate penetration of a prey
animal’s hide and flesh, are rounded both transversely and
anteroposteriorly (Figs. 5B, C). Comparison with the upper
canines of Panthera leo shows that the saber tips of Smilodon
are little, if any, sharper. The force required to drive one, let
alone two, of these formidable looking weapons into a large
prey animal would be enormous.

PALATE

The palate of Smilodon (Fig. 5D) exhibits a pattern of lon-
gitudinal ridges and grooves (Merriam and Stock, 1 932:35—
36). A highly variable medial ridge usually occurs from the
premaxillary-maxillary suture to the posterior margin of the
palatines. In some individuals, this ridge may extend more
anteriorly while in others, parts or all of it may be very
reduced or absent. A lateral ridge occurs on each side of the
medial ridge. These are more consistently developed and
extend posterolaterad from the anterior margins of the an-
terior palatine foramina to terminate between the middle
and posterior of the palatines. The anterior portions of the
lateral ridges bear sharp crests which frequently flare laterally.
Posterior to the palatine-maxillary suture, the lateral ridges
become subdued and broadened before merging with the
palatine surface. A prominent broad groove with a roughened
bottom exists between each lateral ridge and the adjacent
alveolar margin of the palate. Merriam and Stock (1932:36)
suggested that these grooves served as conduits to ingest
sucked blood. This appears to be improbable in that the
posterior portions of the grooves trend laterally and termi-
nate just medial to the M's, not at mid-palate. In addition,
it would be very difficult to seal off the front of the mouth
for effective sucking unless the pool of blood was quite deep
(Bohlin, 1940).

If the mandible is articulated with the cranium in a closed
position (Fig. 5E), however, the rather sharp diastemal crests
align exactly with the middle of the palatal grooves with
about 4 cm of clearance. This alignment of the palatal ridges
and grooves and the diastemal crests of the mandible could
serve as a very effective gripping device for thick pieces of
flesh held in the anterior of the mouth. Galapagos finches
have similar longitudinal ridges and grooves in the homy
palate which line up with the margins of the mandibular
tomia in order to grip and crush seeds (Bowman, 1961). Use
of the palate for gripping would require a much stronger
palatal construction than exists in cats with conical canines.
In fact, the palate of Smilodon is far sturdier than in modem
felids of similar size; the bone is thicker, the corrugations

6 Contributions in Science, Number 356

Akersten: Canine Function in Smilodon

Figure 3. Smilodon. Atlas, LACMHC 2038-5: A. Dorsal view. B. Ventral view. Axis, LACMHC 2039-1: C. Lateral view. Scale bar = 2 cm.
All from Mernam and Stock (1932: pi. 17, figs. 4, 7, 3) with permission, Carnegie Institution of Washington.

would also serve to reinforce this area, and the palate is
partially braced along the midline by the vomer.

CRANIUM-MANDIBLE GEOMETRY

The most important task in determining mode of saber usage
is the investigation of types of motion permitted or excluded
by the geometric relationships of cranium and upper denti-
tion to mandible and lower dentition. The extent of the gape
in Smilodon has often been discussed. I agree with Emerson
and Radinsky (1980) that the maximum possible gape was
about 90 to 95 degrees. As they further point out, this max-
imum gape results in about the same amount of absolute
clearance between the upper and lower canines as does the
maximum gape in modem felids of similar size.

In a detailed study of possible modes of saber use by Smi-
lodon, Simpson (1941) concluded that the momentum need-
ed for a stabbing attack could be generated by an initial leap.
As the sabertooth struck its prey, body momentum would
be transferred to the skull which would whip ventrad to drive
the sabers home. Simpson omitted the mandible from his

figures and calculations, however, stating that inclusion of
the mandible would not have altered his conclusions. If the
mandible is added to all of his diagrams and the outline of
the prey animal’s body is extended in a rounded curve, it is
obvious that the mandible, even at maximum possible gape,
would strike the prey well before or at least simultaneously
with the upper canines. This would either abruptly close the
mouth or prevent canine penetration. Bohlin (1947) also
showed that when the amount of posterodorsad cranial flex-
ion necessary to generate momentum for a stab is considered,
the head would be drawn so far back that the attacker could
not have seen its prey, much less the intended area of attack
(see his fig. 1). Kurten (1952) suggested that sabertooths
pressed the mouth at full gape against the body of their prey
then nodded the head ventrad to stab. While this removes
potential problems which would be created by impact of the
mandible or inability to see the intended prey, there would
now be far too little force for the sabers to penetrate.

Initial penetration of a tapered, sharp blade is best facili-
tated by motion in the direction of its long axis. Any devia-
tion from the axial direction when the target is struck will

Contributions in Science, Number 356

Akersten: Canine Function in Smilodon 7

Figure 4. Smilodon right upper canine, LACMHC 2000-R.9. From left to right: anterior, medial, cross-sectional, lateral, and posterior views.
Scale bar = 2 cm. From Merriam and Stock (1932: pi. 12, fig. 1) with permission, Carnegie Institution of Washington.

Figure 5. Smilodon. A. Reconstruction of major head-depressing muscles (1 = mastoid process, 2 = atlas, 3 = axis, 4 = stemomastoid/
cleidomastoid, 5 = atlantomastoid musculature, 6 = obliquus capitus posterior) drawn by Mark Hallett. B, C. Lateral and posterior stereopairs
of right upper canine tip, LACMHC 2000-R.31. D. Stereopair of palate, LACMHC 2001-2. E. Palatal view of LACMHC 2001-2 with
articulated mandible. Scales: A, bar = 5 cm. B, in mm. C, same as B. D, E, in cm.

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Akersten: Canine Function in Smilodon 9

Figure 6. Geometric relationships of Smilodon cranium and man-
dible based on LACMHC 2001-2. See text for explanation. Drawn
by Mark Hallett.

result in a larger area of contact and require more initial force
to begin a puncture. As noted by Simpson (1941), angulation
of the saber axis during a stabbing motion by Smilodon would
also place a potentially dangerous strain on the saber and its
alveolus. Figure 6 illustrates the long axis of the saber and
other geometric relationships of the skull and mandibles in
Smilodon. Arc A-B represents the axis of the upper canine,
essentially a perfect segment of a circle with its center at C.
The line B-D is tangent to A-B at B and indicates the di-
rection that the tip of the saber must be moving at the mo-
ment it contacts a prey animal if it is to achieve the most
efficient initial penetration. Clockwise rotation about any
point along the radian C-B or its posterodorsal extension
will result in the required instantaneous force along the di-
rection B-D. The posterodorsal extension of C-B extends
through E, the center of rotation for the atlantooccipital ar-
ticulation. Thus, the powerful atlantomastoid musculature
will rotate the head in precisely the proper motion for most

efficient initial penetration of the upper canines. While this
configuration may appear to confirm a stabbing mode of
attack, further geometric considerations show stabbing to be
very improbable, if not impossible. Rotation about point E
would result in the tip of the saber moving through the arc
F-B-G and the tip of the lower incisors, through the arc H-
I. Obviously, the body of a prey animal would have to extend
inside of arc F-B-G to be struck at all by the tip of the saber
but, if the prey’s body extended inside the arc H-I, the lower
incisors or mandible would impact before the sabers. As the
distance between these arcs is less than 3 cm, a very' precise
motion would be required to stab or slash without abruptly
closing the mouth. Even if this could be accomplished, the
curvature of a large prey animal’s body would result in the
axis of the saber being very oblique to its target and only a
glancing blow could be delivered.

If the action of the other head- and neck-depressing mus-
cles were included in a stabbing or slashing action, the center
of the resulting complex motion probably could not be rep-
resented by a single point. The general area about which
movement would center would, however, be farther posteri-
ad and ventrad. This would result in a slightly greater clear-
ance between the paths described by the motions of the tip
of the saber and the anterior end of the mandible. It would
also result in the long axis of the sabers being at an angle to
the direction of force with concomitant loss of efficiency
during initial penetration. In the most extreme case of a
vertical stab (Simpson, 1941: fig. 1A), only about 12 cm of
clearance could be generated. This would still not be enough
for a reasonably safe attack using a stabbing or slashing ac-
tion.

The Rancho La Brea collections of the LACM include at
least 600 fairly complete crania of Smilodon but their sabers
are not as well preserved as the rest of the specimens. The
sabers are usually broken off, shattered, or have slipped out
of the alveoli. In addition, very few crania have associated
lower jaws. As a result, only one cranium with associated
mandible (LACMHC 2001-2) contains an almost complete
saber in original position and can be used to determine ac-
curately the relationships of the upper and lower canines
during closure of the mouth. Manipulation of the specimen
immediately demonstrates that, as the tips of the canines
pass each other, the tip of the lower passes just interior and
anterior to the tip of the upper (Fig. 7B)— exactly the same
mode of canine opposition which allows other mammalian
carnivores to bite with their canines.

The questions of canine opposition and whether or not
sabertoothed mammals could bite with their canines have
been addressed by several earlier workers. Pomel (1843) in-
terpreted wear facets on the lateral surfaces of lower canines
in Felis meganthereon as indicative of canine opposition in
that form. In discussing several Old World taxa of saber-

Figure 7. Associated Smilodon cranium and mandible, LACMHC 2001-2, depicting mouth closing sequence. A. At maximum gape. See
text for explanation of others. Scales in cm.

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Akersten: Canine Function in Smilodon 11

tooths, Fabrini (1890) appeared to believe that canine op-
position occurred in all and hypothesized a mode of attack
in which a bite was used to pierce the body of the prey then
the head was pulled posteriad to shear through the mouthful
of flesh. He also described an upper canine of Machairodus
nestianus which bore a facet on its medial surface formed by
wear against the lower canine. Matthew (1901), however,
denied that canine opposition could have occurred in any
sabertooth. Schaub (1925) noted the presence of canine op-
position in Machaerodus crenatidens but believed that it did
not occur in Machaerodus aphanistis, Machaerodus cultri-
dens, and Smilodon. (Taxonomic designations used above
are those given by the cited authors and may not reflect
currently used synonymies.) Marinelli (1938) specifically
stated that Smdodon was incapable of biting with the canines.
The potential for canine opposition and canine biting in sa-
bertooths appears to have largely disappeared from consid-
eration in subsequent work except for passing mention by
Bohlin (1940) and a comment by Kurten (1963) that the
Smilodontini evidently used their sabers exclusively in stab-
bing, whereas the Homotheriini used them in biting as well
as stabbing and slashing. The retention of canine opposition
in Smilodon, in spite of all the other structural modifications,
strongly indicates that biting with the canines was very im-
portant. If stabbing or slashing were the primary mode of
attack, I would expect canine opposition to have been lost
and a different relationship, more efficiently adapted for stab-
bing or slashing, to have evolved. A similar point was made
by Bohlin (1940) in that the curvature of the sabers was not
suited for a stabbing attack. Can the various adaptations of
Smilodon, then, be explained in terms of a biting mode of
attack?

The arc of lower canine movement centers at the tempo-
romandibular articulation while the center of canine cur-
vature lies several centimeters ventral to this articulation as
previously described (Fig. 6). The non-coincidence of these
centers results in a constantly changing relationship of the
upper and lower canines during closure of the mouth as shown
in Figure 7. As the mouth closes beyond the point where the
tips of the canines pass, the lower canine moves anteriad
relative to the upper or it can be stated that the saber pro-
gressively moves posteriad relative to the mandible, ulti-
mately all but filling the diastema. As a result, the distance
between the tip of the lower canine and the serrate posterior
margin of the upper canine progressively increases. In Figure
7B, the shortest distance between the tip of the lower canine
and the posterior edge of the upper canine is about 10 mm,
in Figure 7C, 30 mm, in Figure 7D, 45 mm, and in Figure
7E, 55 mm. If a piece of flesh were anchored by the tips of
the lower canines (and, perhaps, also by the sharp lower
incisors and the opposing corrugations of the palate and the
sharp diastemal crests of the mandible), closure of the mouth
would result in the upper canines shearing through that piece
of flesh. I propose the term “canine shear-bite” for this type
of bite as opposed to the “canine puncture-bite” used by
living mammalian carnivores.

As is readily obvious from the above photographic se-

quence, normal individuals of Smilodon could not develop
wear facets between the upper and lower canines. However,
only a slight developmental error would result in their con-
tact. One left saber, LACMHC 7037, exhibits wear from the
lower canine on its medial surface (Fig. 9D). The slightly
arcuate facet is longitudinally striated and about 6.5 cm in
total length but only the most distal 4 cm is deep enough to
expose the dentine.

MECHANICS OF THE CANINE SHEAR-BITE

This mode of attack involves more than simple closure of
the mouth with the mandibular musculature. As Smilodon
pressed its gaping mouth against the body of a prey animal
and began to close the mandible, the tips of the opposing
canines started to fold the skin and flesh of the prey (Fig.
8A). By the time that the tips of the canines approached each
other, they produced a strong fold in the anterior region of
the mouth and began to penetrate the “neck” of that fold
(Fig. 8B). At about this stage, the poor lever arm of the
coronoid process and reduced masseteric muculature result-
ed in Smilodon being unable to close its mouth further by
using only the mandibular musculature and the mandible
became nearly stationary relative to the body of the prey. It
was then anchored by the mandibular musculature and by
pressing the well-developed anterior portions of the man-
dibular flanges against the prey. This pair of laterally flared,
narrow ridges would have provided a very good grip when
pressed against an animal, more so than if the entire anterior
of the mandible protruded anteriorly. Completion of the bite
was accomplished by using the head-depressing muscles to
drive the cranium against the immobilized mandible with the
upper canines shearing through the lateral margins of the fold
of flesh (Figs. 8C, D). The serrate distal portion on the an-
terior margin of the saber facilitated initial penetration while
the rounded, non-serrate proximal portion prevented ante-
rior enlargement of the wound at the expense of posterior
shear during later stages of the bite. There may have been a
sequential use of the head-depressing muscles with the at-
lantomastoid musculature, best oriented for initial saber pen-
etration, acting first. Those which move the axis of the saber
more obliquely would, perhaps, contract in later bite stages.
The short lower canines, with probably some assistance from
the tips of the lower incisors, could only penetrate deeply
enough to anchor the fold. At some intermediate stage, the
opposing ridges and grooves of the palate and the diastemal
ridges of the mandible gripped and helped to stabilize the
fold of flesh. As the head was depressed, the posterior of the
mandible would also move ventrad and rotate the anterior
so that more of the mandibular flange would contact the prey.
This anchored the mandible even more firmly to oppose head
depression and resulted in the incisors rotating away from
the prey to stretch and thin the neck of the flesh fold. During
the final stages of the bite, the upper and lower incisor bat-
teries (which now functionally included the lower canines)
punctured the neck of the flesh fold with a closely spaced
series of perforations. Smilodon could then free the entire

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Akersten: Canine Function in Smilodon

Figure 8. Reconstruction of canine shear-bite sequence in Smilodon. See text for explanation. Drawn by Mark Hallett.

fold of flesh with only a modest pull, leaving a major wound
in the body of its prey.

SUPPOSED EVIDENCE OF STABBING FROM
WOUNDS FOUND IN SKULLS

Three specimens have been described as bearing wounds
attributed to stabbing by sabertoothed mammals. Scott and
Jepson ( 1 936) described a skull of Nimravus with an oblique-
ly directed, elongate, and partially healed wound to the left
frontal. They noted that the size and shape of the wound
conformed to the upper canine of Eusmilus and concluded
that this specimen provided evidence that Eusmilus stabbed

with its sabers. The size and shape of the injury appear to
be consistent with damage inflicted by a saber, but the spec-
imen lacks conclusive evidence— part of a saber within the
wound. Even if we assume that a saber caused the damage,
it need not have resulted from a stab. A saber puncture could
have just as easily, or more easily, been accomplished with
a bite. The head of Nimravus appears to be small enough to
have been taken into the mouth of Eusmilus and bitten. This
specimen does not provide any solid evidence regarding the
mode of attack used by sabertooths.

Miller (1969, 1983) used the above specimen and two
others from Rancho La Brea to support a stabbing attack for

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Akersten: Canine Function in Smilodon 13

Figure 9. Varanus komodoensis, LACM Herpetological Collections 121971. A. Stereopair of upper tooth. C. Upper dentition. Damage to
Smilodon scapulae inflicted by a square-ended pick about 1915. B. LACMHC K-50, stereopair of pick entry hole. E. LACMHC K-232,
stereopair of oblique blow from pick. F. LACMHC K-50, reverse side of different pick hole from that shown in B. Note depressed bone flake,
lack of bone flake detachment, and splintery fractures usually typical of damage to fresh bone. D. Stereopair of medial surface of Smilodon
left upper canine showing wear facet from lower canine. Scales: A and B in mm, others in cm. B and F coated with ammonium chloride.

sabertooths. The only information available for one of the
latter specimens is a brief mention by Moodie (1923:128)
“A skull of a young wolf the brain case of which is cut through
by the tooth of a tiger, the saber being broken off and imbed-
ded in the preserved skull, is on exhibition at the University
of California.” Since this specimen has never been illustrated
or adequately described and cannot presently be located in
the Berkeley collections (J.H. Hutchison, pers. comm., 1983),
it does not provide useful information about mode of saber
use. The other La Brea specimen cited by Miller (1983) is a
Smilodon skull (LACMHC 2001-24) with an elongate,
anteroposteriorly oriented perforation of the left frontal. Rel-
ative to the skull orientation, this injury has a rounded pos-
terior margin and a pointed anterior margin, very similar to
the cross section of a saber oriented so that the anterior
margin of the saber is at the posterior margin of the perfo-
ration. An isolated saber, however, can be inserted from
either direction. S. Shermis (pers. comm., 1983) conducted
a pathology study of this specimen and concluded that char-
acteristics of the injury are compatible with insertion of a
saber from the anterior of the individual while the animal

was still alive. In Rancho La Brea specimens, however, dam-
age known to be of recent origin often has characteristics
considered to be indicative of injury which occurred during
life or soon after death (Figs. 9B, E, F). This probably results
from the unusually good preservation of collagen in bones
from Rancho La Brea (Ho, 1965; Doberenz and Wyckoff,
1967).

While the conclusive evidence of a broken saber in the
wound is again lacking, the evidence does indicate that an
upper canine of Smilodon could have caused the damage.
But is this evidence for stabbing? With a maximum 3-cm
clearance between the arcs described by the tip of the canine
and the tip of the mandible, the position of the injury does
not allow sufficient clearance for either an anterior or pos-
terior attack without the attacker first striking its mandible
against the skull. In order to determine if any method of
stabbing would be feasible, the injured specimen was ma-
nipulated with another skull bearing mandibles mounted at
maximum gape. All four attack possibilities were attempted:
posterior stab with left saber, posterior stab with right saber,
anterior stab with left saber, and anterior stab with right

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Akersten: Canine Function in Smilodon

saber. In both the posterior stab with left saber and the an-
terior stab with right saber, the opposite saber contacted the
skull in the area mediad to the right postorbital process and
prevented insertion of the attacking saber. In both posterior
attacks, contact of the mandible with the posterior of the
injured cranium prevented the attacking saber from striking
the skull at the proper angle to produce the observed injury,
but a more elongate perforation could have been produced.
Both types of frontal attacks had similar results from the
mandible striking the anterior of the head. The observed
injury could not have been the result of an open-mouthed
stab by another Smilodon. It could, however, have been the
result of a canine blow struck with the mouth closed. An
upper canine of Smilodon inserted in the injury does pene-
trate to the depth consistent with the maximum length of
exposure ventral to a closed mandible. Therefore, it seems
plausible that (if one assumes that a saber was actually re-
sponsible for the perforation) the injury may have been caused
by an accidental blow struck with the mouth closed during
intraspecific combat.

A specimen recently collected by the Rancho La Brea Proj-
ect deserves mention here. The left third rib of a Smilodon
(LACMRLP R25876) had the very tip of a Smilodon upper
canine (separately cataloged as LACMRLP R25877) imbed-
ded in its anterolateral surface. This specimen was found
with the left fourth rib of a Smilodon in very close to life
position but no other elements can be reasonably associated
with the two ribs. The puncture produced by the canine
appears to have been made in green bone. Its long axis is
nearly transverse to the length of the rib with the anterior
end (relative to rib orientation) slightly dorsad to the pos-
terior end. The saber tip is obliquely broken; the distance
from the tip to the break is between 3 and 6 mm. The margin
with the greatest portion remaining appears to be the pos-
terior and occurred in the posterior margin of the puncture.
Therefore, the bite was probably made from a posterior and
slightly ventrad orientation. Depending on the forelimb ori-
entation of the bitten individual, the saber may have passed
through the posterior part of the scapula. None of the Smi-
lodon left scapulae from this area of the excavation exhibits
possible saber damage but several lack the posterior portion.
Other specimens from Rancho La Brea also suggest intra-
specific strife (S. Shermis, pers. comm., 1983).

It is interesting that all of the specimens which supposedly
show some evidence of damage inflicted by the upper canines
of sabertooths are from carnivores. The lack of herbivore
specimens displaying saber injury is indirect evidence that
bony areas were avoided during predation. Intraspecific or
interspecific combat among carnivores is quite different from
predatory attack behavior (e.g., Schaffer, 1972; Bertram, 1978;
Leyhausen, 1979). Therefore, even if the wounds in these
specimens were actually caused by sabers during life, they
should not be considered as indicative of sabertooth pred-
atory behavior.

SOCIAL STRUCTURE IN SMILODON

Hunting techniques employed by various predators are de-
pendent in part upon whether they seek prey in a group with

cooperative behavior or alone (except for a female with young).
Although this is almost impossible to interpret for extinct
forms which do not have close living relatives, a few infer-
ences can be drawn about social structure in Smilodon from
the fossil record at Rancho La Brea. As Gonyea (1976) point-
ed out, the large numbers of Smilodon preserved at this
locality strongly suggest that they lived in groups or prides.

The LACM made a large collection of fossil vertebrates,
now termed the Hancock Collection, from Rancho La Brea
between 1913 and 1915. Few of these specimens have been
exchanged or lost so that the relative numbers of the larger
taxa presently in this collection should be a very close ap-
proximation to the preserved thanatocenose. Only about one
third of the Hancock Collection, roughly 260,000 specimens,
has been cataloged to date. A number of years will be required
to complete the task and produce a definitive census of the
megafauna, but it is obvious that larger taxa with the lowest
proportion of cataloged elements are coyote and Smilodon.
Marcus (1960) used the catalogs plus complete counts of
Bison and Camelops in his census of the megafauna and
arrived at a total of 1029 as the minimum number of Smi-
lodon in the collection. Miller (1968), however, estimated
about 2100 individuals of Smilodon based on cranial ele-
ments.

Unpublished data, briefly reviewed by Akersten, Shaw,
and Jefferson (1983), from a recent excavation at Rancho La
Brea indicate that entrapment in shallow asphalt seepages
was the primary mode of producing the rich fossil deposits
at Rancho La Brea. An episode of entrapment apparently
began when one large animal (probably an herbivore since
these would be most common in natural large njammal pop-
ulations) blundered into a shallow puddle of asphalt. The
helpless or dead herbivore would then attract a number of
opportunistic carnivores, thus accounting for the fact that
carnivores make up about 90% of the larger mammals found
at this locality. The total number of large herbivores should
closely approximate the total number of entrapment epi-
sodes. Marcus (1960) counted 423 individuals of large her-
bivores (Bison, Camelops, Nothrotheriops, Glossotherium, and
Equus). Allowing for a few proboscidians and uncataloged
or lost specimens, the maximum number of individual her-
bivores and, consequently, the number of entrapment epi-
sodes represented in the Hancock Collection is considerably
fewer than 600. Therefore, an average of 1 .7 to 3.5 Smilodon
were caught during each entrapment episode, depending on
the count used for Smilodon. As it is unlikely that all en-
trapped herbivores lured Smilodon in equal numbers, as many
as six or eight may have been caught during a single entrap-
ment episode — a period of weeks or several months at the
most. This ratio would be improbable if Smilodon were a
solitary hunter unless, unlike living large predators, a number
of individuals shared overlapping hunting areas. The best
explanation of these data, providing that the mode of en-
trapment is correctly interpreted, is that Smilodon was a
social animal and may have hunted in groups.

Another way of looking at the same data is to compare
the relative numbers of Smilodon at Rancho La Brea with
those of other predators, whose social structure is known or

Contributions in Science, Number 356

Akersten: Canine Function in Smilodon 15

can reasonably be inferred. The other very common predator
is Canis dims Leidy with at least 1646 individuals repre-
sented in the Hancock Collection (Marcus, 1960), approxi-
mately equal to Smilodon. Though C. dims is extinct, its
morphology is quite similar to the extant C. lupus and a
reasonable conclusion would be that it also hunted in fairly
large groups. The other common, but smaller, predator is the
extinct C. latrans orcutli (Merriam), very closely related to
the living coyote (Nowak, 1979). Marcus accounted for 239
coyotes in the Hancock Collection but, as previously noted,
much of the coyote material has not been cataloged. Coyotes
are also social animals but usually form smaller groups than
wolves. Carnivores which do not hunt in packs, Fe/is con-
color, Panthera onca (Jefferson, 1983), Ursus americanus,
Ursus arctos, Lynx rufus, Urocyon cinereoargenteus, and
Taxidea taxus, are comparatively rare at Rancho La Brea.
Thus, if the relative abundance of carnivores preserved at
Rancho La Brea reflects hunting behavior, Smilodon must
be included among the social forms. As the American lion,
Panthera atrox (Leidy), is relatively uncommon with 76 in-
dividuals reported by Marcus, the same line of reasoning
suggests that this predator was either a solitary hunter or, if
it hunted cooperatively, groups did not often frequent the
area.

Finally, evidence that Smilodon may have been a social
animal derives from the relatively high frequency of indi-
viduals from Rancho La Brea which either could not have
killed prey or would have had great difficulty in doing so.
Seven adult skulls in the collection had only one saber during
life as shown by the undeveloped or secondarily lost alveolus
for the other. The remaining canine alveolus in one of these
specimens is small and distorted. One or both canines in
several skulls were broken off and subsequently worn during
life. Individuals lacking one or both canines would probably
experience difficulty making a kill. Many postcranial ele-
ments of Smilodon in the Hancock Collection exhibit pa-
thology, some very severe (Moodie, 1926, 1927, 1930). At
least one limb was all but useless in some individuals; ver-
tebral abscesses and fusions would have hampered or crip-
pled many more. A number of skulls also have badly worn
cheek teeth (Miller, 1968). Solitary carnivores possessing such
major disabilities soon perish unless they manage to survive
by scavenging. These same disabilities, however, would place
them at an extreme disadvantage when competing with
healthy individuals for carrion. Most living large carnivores
regularly attempt to appropriate carcasses from other pred-
ators. Social carnivore groups, on the other hand, frequently
allow incapacitated individuals to feed on kills made by other
pack members. Schaller (1972) and Bertram (1978) describe
a number of occasions when African lions (especially females)
incapacitated by age or injury, survive by feeding on kills
made by other pride members. Even aged nomadic males
may survive as members of nomadic groups or by being
allowed to feed at the kills of other nomads (e.g., Schaller,
1972:81). Schaller (1972:358) even considers that one func-
tion of the lion’s social system is to provide “life insurance”
for individuals unable to hunt for themselves but the selective
advantage of this is difficult to envision. Kruuk ( 1 972) noted

that older female members of spotted hyaena clans, no longer
able to run well, feed from kills made by other clan members.
The African wild dog appears to have the most highly de-
veloped social structure among the fissiped carnivores. Estes
and Goddard (1967:68) report that the pack provides food
for “sick and old adults unable to kill for themselves.”

It has been suggested that the frequency of pathologic Smi-
lodon specimens from Rancho La Brea was a result of crip-
pled individuals specializing in feeding from the carcasses of
trapped animals (Bohlin, 1947). However, if one assumes
that the Hancock Collection represents only 10% of the total
individuals of all species that were once trapped in these
deposits, the 600 episodes of entrapment represented by this
collection become 6000, spread over at least 25,000 years or
an average of no more than one every four years. Even al-
lowing for a probable clustering of entrapment episodes
through time, this could hardly represent a dependable source
of food, especially considering the hazards of scavenging at
such a place and the necessity of competing with healthy
carnivores for carrion.

Although the data regarding Smilodon' s social behavior
are far from conclusive, they are more easily explained by a
cooperative model than by a solitary model. It is not possible,
however, to draw inferences about the extent of cooperation
during the hunt. They may have merely hunted the same
area with each attack made by an individual acting on its
own, then fed as a group, or they may have cooperated to a
greater degree throughout the hunt.

PREY SPECIES

It has generally been assumed that the sabers of Smilodon
and other sabertooths were adaptations for attacking large,
relatively thick skinned prey such as ground sloths or pro-
boscidians. The only direct evidence available comes from
the Late Pleistocene fauna of Friesenhahn Cave in Texas
(Evans, 1961; Meade, 1961; Lundelius and Slaughter, 1971;
Graham, 1976, pers. comm., 1983; Rawn-Schatzinger, 1983).
This cave appears to have been a denning site for Homo-
therium to judge from the number of individuals recovered
and especially from the occurrence of several very juvenile
specimens. Scores ofjuvenile proboscidians, primarily mam-
moths, were also found but adult proboscidians and other
large carnivores are rare. This association of an apparent
denning site with the remains of juvenile proboscidians
strongly suggests that Homotherium preferentially hunted
juvenile proboscidians and brought their remains back to its
lair. Perhaps the coarsely serrate margins of Homotherium ’s
sabers and other teeth facilitated dismembering the carcasses
into more easily carried chunks. The incisors, in particular,
are almost shark-like with very serrate margins.

If the herd and parental behavior of extinct proboscidians
were similar to those of modem African elephants, hunting
juveniles could have been a hazardous occupation. Many or
all of a herd of elephants will defend or aid any juvenile; they
may even continue to defend the body several days after
death (Douglas-Hamilton and Douglas-Hamilton, 1975).
Even if mammoth behavior were more like that of Indian

16 Contributions in Science, Number 356

Akersten: Canine Function in Smilodon

elephants with separate nursing and juvenile care units, as
suggested by Graham (1976, pers. comm., 1983), attacking
predators would have had to be cautious. Whatever the case,
Homotherium probably waited for an ideal opportunity and
retreated immediately after even a successful attack on a
juvenile until the adults left the area.

It is certainly tempting to extend this interpretation of
Homotherium ’s prey to Smilodon (and other sabertooths),
but the two genera have different morphologies (Meade, 1961;
Churcher, 1966), even different dental eruption sequences
(compare Rawn-Schatzinger, 1983, with Tejada-Flores and
Shaw, 1984). In addition, the shear-bite of Smilodon would
have been equally effective on smaller, thin-skinned prey.
The powerful build of Smilodon indicates that they probably
could have successfully attacked any of the larger herbivores
found at Rancho La Brea except for adult proboscidians.
They were certainly attracted to entrapped non-proboscidi-
ans because they are quite common in many La Brea deposits
that totally lack proboscidian remains. I doubt that Smilodon
fed exclusively on members of any one taxon — no living large
predator does— but they may have more commonly hunted
juvenile proboscidians than did the other Carnivora found
at Rancho La Brea.

AREA OF ATTACK

Most researchers have concluded that the upper canines of
sabertooths were too fragile to be used on bony areas of their
prey but Gonyea (1976), rebutted by Emerson and Radinsky
(1980), thought that a stab at the back of the neck or skull
was more likely. Even though the sabers of Smilodon are
much heavier than a knife, they still have a rather slender
cross section in comparison to their length. Because of their
length, it seems likely that relatively little lateral force near
the tip would cause breakage. If they were used in an attack
on a bony area, one saber would probably contact bone before
the other, resulting in considerable lateral torque and prob-
able breakage (Bohlin, 1947). Repeated contact with bone
would also cause wear on the tips of the canines. Leyhausen
(1979:33) notes that even the much stouter conical canines,
with thicker enamel, of modem cats readily splinter from
normal use.

The curated portion of the Hancock Collection was sur-
veyed for sabers with well-preserved tips and fully formed
roots. Of 54 adult sabers, one displayed a minute wear facet
on the enamel of the tip, two had wear that barely penetrated
to the dentine, and only one had an appreciably wear facet:
2 mm wide by 2.5 mm long. Three others exhibited breakage
of the very tip with some subsequent wear; this suggests that
the one specimen with a larger wear facet may have also
resulted from wear after breakage. All of the sabers with tip
wear are isolated specimens exhibiting moderate to extreme
wear of the serrations and appear to come from older indi-
viduals. This information supports the interpretation that
the sabers were not employed in attacks on bony areas. The
sabers with broken and worn tips, the occurrence of a saber
tip in a Smilodon rib, and the skulls with more severely
broken sabers showing post-breakage wear do show that mis-

takes were occasionally made. Of the 17 juvenile sabers lo-
cated, 10 had appreciable wear facets on the tips and one
had a minute facet. The larger facets tended to be oblique
with more wear toward the lateral margins. Perhaps Smi-
lodon kittens were less careful with their sabers than adults.

If the interpretation is correct that the canine shear-bite of
Smilodon was normally directed toward areas in which bone
would not be encountered, only the throat and abdomen are
possible targets. The throat has been suggested as the focus
for a stabbing or slashing attack because of the shallow carotid
artery and jugular vein and because most modern felids typ-
ically employ a nape or throat bite (Martin, 1980; Emerson
and Radinsky, 1 980). It seems to me that a throat bite would
have to be delivered with precision in order to sever these
blood vessels without encountering the cervical vertebrae or,
in short-necked juvenile proboscidians, without striking the
posterior of the mandible or the anterior of the humerus.
Because the tips of the sabers are well outside the visual area
of Smilodon, a throat bite would be potentially hazardous
to these teeth. Leyhausen ( 1979) states that the neck bite in
living felids results from a taxis oriented toward an inden-
tation (the neck) between a large cylinder (the body) and a
smaller one (the head). This taxis does not discriminate be-
tween throat and nape orientations; the nape bite used by
smaller felids is learned. Larger felids may use either nape
or throat bites (other points and methods of attack less com-
monly), possibly depending on prey size or on the learned
behavior of the individual. One interesting variation is the
bite to the posterior of the cranium employed by jaguars in
killing capybaras (Schaller and Vasconcelos, 1978). A jugu-
lar/carotid attack by Smilodon would require a much more
specific taxis than occurs in modem felids, because the pro-
cess of learning the exact point to bite by trial-and-error
would be far too hazardous to the sabers. I also find it difficult
to explain the development of extremely elongate sabers in
Smilodon and other dirktooths in terms of a throat attack.
The shorter sabers of scimitartooths should be more than
sufficient to sever the major blood vessels.

The elongate sabers of Smilodon appear to be adapted for
causing massive damage with a single bite. The large, bone-
free abdominal region, with a rich supply of blood vessels
and a variety of vital organs, would be a more logical area
to wield these weapons. Furthermore, unlike the neck, there
would be relatively little muscle tissue for the incisors to
penetrate at the end of the bite. The type of accuracy needed
in order to sever the jugular/carotid blood vessels without
striking bone would not be required, simply a taxis directed
toward the region anterior to the hind legs. Most authorities
agree that, while large, living felids rarely attack the abdo-
men, they do typically begin feeding there (e.g., Leyhausen,
1979; Schaller, 1972). African lions, after pulling down large
prey such as rhinoceros and hippopotamus, are known to
occasionally attack the abdomen instead of the throat (J.
Kingdom pers. comm., 1983). Wolves (Young, 1944; Mech,
1970), African wild dogs, golden jackals (van Lawick-Good-
all and van Lawick-Goodall, 1971), and spotted hyaenas
(Kruuk, 1 972) frequently kill by attacking the abdomen. This
is probably the only vulnerable area easily available to such

Contributions in Science, Number 356

Akersten: Canine Function in Smilodon 17

Figure 10. Reconstruction of initial stages of attack by Smilodon. Left, pulling down young mammoth. Right, beginning of canine shear-
bite to abdomen. Drawn by Mark Hallett.

packs of relatively small predators which lack the claws and
bulk to pull down their prey in order to attack other vita!
areas. Even so, it does show that abdominal attack can be
effectively used by modem mammalian carnivores.

Martin (1980) considered an abdominal attack by saber-
toothed cats, especially by dirktoothed forms such as Smi-
lodon, to be very improbable. He argued that the prey would
not be killed immediately and, unless it went into total shock,
would try to escape from the attacker which was not adapted
to pursue an escaping animal. He also claimed that the ab-
domen can be defended by the head of the prey and that
stabbing a broad, gently sloping abdominal area would be
difficult at best. I concur with the last point; however, the
canine shear-bite is perfectly adapted for attacking such areas.
Furthermore, it is difficult to visualize how the forms which
were potential prey for Smilodon at Rancho La Brea could
have used their heads to defend their abdomens while
stretched out on the ground. Figure 10 depicts Smilodon
pulling down a juvenile mammoth and initiating a canine
shear-bite to its abdomen.

in regard to the prey attempting to escape, Schaller (1972:
266) stated that prey pulled down by lions and not yet bitten
appeared to go into shock and rarely struggled to any extent.
He went on to describe an uninjured buffalo that lay on its
side while its tail was chewed by a lioness. Auffenberg (1981)
credited shock as being important in the lack of struggle
evinced by downed but not yet killed prey of Varanus ko-
modoensis. Furthermore, tigers are easily able to control large,
struggling prey after bringing them down (Schaller, 1967). I
doubt that prey would make much of an attempt to escape
after the shock of being pulled down by Smilodon and having
a huge chunk of the abdominal region torn out. The claws

and powerful forelimbs of Smilodon would easily be able to
control any who might try to struggle. Even if an occasional
prey animal could escape after the attack, its severe injuries
would prevent it from running very fast or very far.

A POSSIBLE KILL SCENARIO

Events in a typical attack sequence might have taken the
following course, assuming that Smilodon did cooperate to
a minor extent during the hunt, that the prey in this particular
case was a juvenile mammoth, and that mammoth social
behavior was similar to that of modem Indian elephants
(Graham, 1976, pers. comm., 1983).

A pride of sabertooths scatter out while approaching a herd
of adult female and juvenile mammoths, targeting one pair
of juveniles who were playing a short distance from the rest.
While several of the predators distract the herd, one makes
a short rush from concealment and pulls one juvenile down
toward itself with the retractile claws and powerful forelimbs.
Quickly orienting itself to the posterior of the abdomen, the
sabertooth opens a gaping wound with a canine shear-bite,
then flees before the mother and the rest of the herd can
retaliate. The pride regroups at a distance and waits for the
critically injured juvenile to die and for the rest of the herd
to leave. Once the dead animal is abandoned, the sabertooths
(including aged or incapacitated members of the pride) return
to feed. They tear the carcass apart with their prognathus
incisors and occasionally employ a canine shear-bite to open
up a new area. The rather long lips of sabertooths allow them
to take chunks and strips of flesh into the side of the mouth
so that the highly developed camassials can slice the meat

18 Contributions in Science, Number 356

Akersten: Canine Function in Smilodon

into pieces small enough to swallow (Miller, 1969; Martin,
1980).

A MODERN ANALOG, THE KOMODO DRAGON

A major problem in interpreting the mode of attack in sa-
bertooths has always been the lack of a modern analog. No
living mammalian predator has teeth comparable to the sa-
bertooth canine and none is known to consistently kill its
prey by first pulling it down then biting open the abdomen,
as I hypothesize for Smilodon. If one looks at non-mam-
malian predators, however, one very interesting modern rep-
tilian analog stands out: the Komodo dragon ( Varanus ko-
modoensis). Auffenberg (1978, 1981) has thoroughly studied
this large active predator, which may grow up to 3 m long
and 60 kg in weight. Its dentition consists of mediolaterally
flattened, sharp, recurved teeth with serrations on the entire
posterior margins and about the distal one fourth of the
anterior margins. Although this reptile has numerous teeth
in both upper and lower dentitions, the individual teeth quite
closely resemble the canines of Smdodon, even in the dis-
tribution of serrations (Figs. 9A, C).

The usual prey of the Komodo dragon consists of deer and
boar. Recorded kills include deer of up to 80 kg and boar up
to 40 kg, but villagers report kills of deer up to 200 kg. The
Komodo dragon typically kills these prey by ambush along
game trails or in bedding areas. Because it lacks prehensile
forelimbs, it grasps them with the mouth, pulls or wrestles
them down, then bites open the abdomen. Small individuals
may be picked up in the powerful jaws and shaken. Attacks
on tethered goats indicate that they appear to be in shock
prior to their abdomens being ripped open; death probably
results from “massive viscaral bleeding” (Auffenberg, 1981:
247). As Auffenberg further notes, visceral bleeding could be
enhanced by physiological shock resulting from the violent
attack.

Successful attacks on larger prey, such as tethered or free
water buffalo up to 590 kg, apparently follow a different initial
pattern. The Komodo dragon lacks the strength to bring down
such large prey but a few individuals learn to repeatedly bite
and slash at the legs until the animal is crippled by the sev-
ering of its tendons and collapses. Available indirect evidence
indicates that the kill is again accomplished by biting open
the abdomen (Auffenberg, 1981:261). Unlike Smilodon, the
Komodo dragon can afford to attack bony areas such as lower
legs because it has numerous and replaceable teeth; however,
it still makes the killing bite to the soft abdomen after bring-
ing the animal down. That it can easily bite open the ab-
domen of a large water buffalo with saber-like teeth only 2
cm or less in height surely shows that the sabers of Smilodon
could very effectively function in a similar manner. The
method of biting also differs in that the Komodo dragon
delivers repeated bites with backward jerks at the same spot.
Despite the number of differences, I suggest that this reptile
is the closest living analog to Smilodon in kill methodology.

Other indirect evidence tends to support the analogy. The
prey animals of the extant Komodo dragons appear to have
been introduced by man. Pleistocene deposits on the island

of Flores (within the present range of the Komodo dragon)
appear to contain only two large animals, both miniature
stegodont elephants (Hooijer, 1972a). On Timor, east of the
Komodo dragon’s present range. Pleistocene deposits have
yielded large varanid vertebrae similar to Varanus komo-
doensis in association with the same proboscidians and giant
tortoises (Hooijer, 1972b). Large varanid vertebrae are also
known from the Pleistocene of Java (Hooijer, 1972b) with a
more varied fauna including proboscidians. These data led
Auffenberg (1981:289) to suggest that the ancestors of Ko-
modo dragons once fed on small proboscidians. Thus, a ten-
tative parallel can be drawn between the prey of Pleistocene
Komodo dragons and, at least, Homotherium.

In addition, Australia stands out as the only temperate
continent which lacks the remains of some type of saber-
toothed mammal. This seems odd because the carnivorous
marsupials there underwent as extensive a radiation as did
the South American marsupials which did evolve a saber-
toothed form. Varamds are, however, known in Australian
faunas by the Middle or Late Miocene and underwent a major
radiation, culminating in the giant. Late Pleistocene Mega-
lania prisca Owen which reached a total length of perhaps 7
m and a maximum weight of 600 to 620 kg (Hecht, 1975).
The teeth of Megalania closely resemble (but are larger than)
those of Varanus komodoensis, suggesting a similar, highly
predaceous mode of life. Hecht believed that Megalania was
the major predator of the Late Pleistocene giant marsupials
of Australia. Perhaps Megalania and other large fossil Aus-
tralian varamds occupied a niche similar enough to saber-
toothed mammals that they precluded marsupial carnivores
from that adaptive zone.

CONCLUSIONS

The hypothesis that Smilodon pulled down its prey, then
killed with a canine shear-bite to the abdomen, appears to
be consistent with all of the observed morphology relevant
to use of the upper canines and eliminates several anomalies
introduced by stabbing or slashing hypotheses. The powerful
head-depressing muscles function to bite by means of de-
pressing the cranium against an essentially stationary man-
dible, held in place by pressing the anterior margins of the
mandibular flanges against the prey. The mandible needs to
be very robust to resist the developed forces but does not
require a long coronoid process in order to bite with the
canines. The lower canines are relatively small because they
need only penetrate enough to anchor the fold of flesh taken
into the mouth until the very end of the bite, then they
function with the incisor battery. In the shear-bite model, a
tremendous force is no longer needed to drive the sabers into
the prey. The geometric relationships of the cranium, man-
dible, and dentition are far easier to explain. The palatal
ridges and grooves serve as gripping devices in the absence
of most of the premolars. One could even speculate that the
retracted nose of Smilodon (Miller, 1969) was an adaptation
to avoid friction bums from rubbing against the hair of its
prey during the head depressing stages of the canine shear-
bite. The problem of structural and behavioral intermediates

Contributions in Science, Number 356

Akersten: Canine Function in Smilodon 19

between biting and stabbing/slashing forms disappears. The
similarities between the kill behavior hypothesized here for
Smilodon and that observed in the living Varanus komo-
doensis, plus the correspondence in dental morphologies be-
tween the two, add the dimension of a modem analog.

Examination of fairly complete and undistorted speci-
mens, casts, and figures of other sabertoothed mammals in-
dicates that they all very probably possessed upper and lower
canine occlusion similar to that of Smilodon. Therefore, I
believe that the canine shear-bite was utilized by all, even
though the details of its use must have differed as demon-
strated by the variety of other morphologic features exhibited
by various taxa. The shorter sabers of Homotherium may
have been more useful in a throat attack and the coarser
nature of their serrations may indicate that these teeth were
more frequently employed for some purpose (such as to dis-
member carcasses) in addition to the kill. In Thylacosmilus,
the lack of upper incisors, the ever-growing upper canines,
the blunt lower canines which honed the uppers, and the
longer series of cheek teeth (Riggs, 1934; Turnbull, 1978)
clearly show that the canine shear-bite of this genus must
have differed in many details. The morphological differences
between other sabertooths and Smilodon are many, but I
believe that they do not negate the hypothesis that all em-
ployed some variation of the canine shear-bite to kill prey.

ACKNOWLEDGMENTS

I dedicate this paper to the memory of my father, Henry
Akersten, who taught me the joys of following one’s curiosity.
Most of this study was conducted while on the staff of the
George C. Page Museum. Many individuals contributed sug-
gestions and comments during the evolution of this concept.
Among them I especially thank C.A. Shaw of the George C.
Page Museum, L.F. Marcus of Queens College, G.J. Miller
of the Imperial Valley College Museum, and W.D. Turnbull
of the Field Museum. The 1980 Society of Vertebrate Pa-
leontology banquet address by W. Auffenberg, University of
Florida, on the Komodo dragon brought it all together. W.
Langston, Jr., of the Texas Memorial Museum provided casts
of Homotherium and J.W. Wright and R.L. Bezy of the Nat-
ural History Museum of Los Angeles County made available
the specimen of Varanus komodoensis. C.A. Shaw and J.M.
Harris reviewed and greatly improved early drafts of the
manuscript. Drawings reprinted from Merriam and Stock
(1932) were prepared by John L. Ridgway. The remaining
drawings were prepared and generously donated by Mark
Hallett. Richard Meier and John DeLeon assisted with pho-
tography. I especially thank Mr. Meier for developing the
double exposure technique used for Figure 1G. Olive and
Antonia fejacia- Flores provided invaluable translation as-
sistance. Specimens with an LACMRLP prefix were collected
and curated with the aid of grants from the Natural History
Museum of Los Angeles County Foundation and the Na-
tional Science Foundation (NSF GB 24819). Other speci-
mens came to light as a result of a National Science Foun-

dation Biological Research Resources Program grant (NSF

BSR 82-18194).

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Contributions in Science, Number 356

Akersten: Canine Function in Smilodon 21

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Submitted 1 1 November 1983; revised and accepted 26 July
1984.

22 Contributions in Science, Number 356

Akersten: Canine Function in Smilodon

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S Y STEM ATICS AND DISTRIBUTION OF THE SKINKS
ALLIED TO EUMECES TETRAGRAMMUS
(SAURIA: SCINCIDAE)

Carl S. Lieb

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Natural History Museum of Los Angeles County * 900 Exposition Boulevard » Los Angeles, California 90007

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SYSTEMATIC'S AND DISTRIBUTION OF THE SKINKS
ALLIED TO EUMECES TETRAGRAMMUS
(SAURIA: SCINCIDAE)

Car! S. Lie!)1

ABSTRACT. The distributions and relationships of the three species
in the Eumeces brevilineatus Group, E. brevilineatus, E. calliceph-
alus, and E. tetragrammus are re-evaluated. E. brevilineatus and E.
tetragrammus are sympatric over a narrow zone in southern Texas,
and intermediates in color pattern occur at some localities. The
presence of intermediates and lack of other morphological di fferences
between the two taxa suggest that gene exchange occurs in the area
of contact.

The third member of the species group, callicephalus, is the most
distinctive in scalation and color pattern. It is allopatric to the dis-
tributions of brevilineatus and tetragrammus, but populations of
brevilineatus that are geographically nearest to the range of calliceph-
alus contain some individuals with the distinctive traits of the latter
form. The expression of these characteristics within the westernmost
populations of brevilineatus is interpreted as evidence of former gene
flow that was interrupted by Pleistocene alterations in the habitat
and climate of the western Chihuahuan Desert.

The three nominal species of the E. brevilineatus Group are rel-
egated to subspecies of tetragrammus (Baird). Salient features of
coloration and scalation of the polytypic tetragrammus ally it with
anthracinus and septentrionalis of the E. anthracinus Group. A total
of 13 species groups in the genus are now recognized; a key to these
groups and a list of the constituent species are provided.

RESUMEN. En su monografia del genero Eumeces, Taylor ( 1 935b)
asigno tres especies de lagartijo escincidos a La Especie-Grupo Eu-
meces brevilineatus: E. brevilineatus de la parte norte de Mexico y
Tejas, E. callicephalus de la parte oeste de Mexico y sur de Arizona,
y E. tetragrammus de la parte nordeste de Mexico y sur de Tejas.
Las especies E. brevilineatus y E. tetragrammus fueron clasificadas
por diferencias en deseno de las coloracion, poseiendo E. brevilin-
eatus bandas y rayas sobre el cuerpo mas cortas que E. tetragram-
mus. Aun cuando las dos son escencialmente identicas en sus es-
camas, Taylor las clasifico como especies diferentes porque el no
encontro invididuos intermedios, y porque supuso la existencia de
una area grande simpatrica en sus distribuciones geograficas. Sin
embargo, una revista de las distribuciones de estas dos especies revela
que su area de distribucion geografica simpatrica esta confinada a
una zona estrecha en el sur de Tejas. Ademas, el examen de espe-
cimenes obtenidos hasta el presente, indica que rangos intermedios
en coloracion existen, y por eso, es posible que las dos formas in-
tercambien material genetico en la area de contacto.

El tercer miembro de las especie-grupo, Eumeces callicephalus,
tiene una distribucion geografica que es alopatrica a las distribuciones
de E. brevilineatus y E. tetragrammus. E. callicephalus se localiza
en elevaciones bajas y moderaciones al oeste de la Division Conti-
nental, desde la zona central de Jalisco hasta el Sur de Arizona. De
los tres miembros del Grupo E. brevilineatus, E. callicephalus es el
mas distinto en sus escamas y coloracion. Empero, poblaciones de
E. brevilineatus que se encuentran mas proximas en su posicion
geografica a poblaciones de E. callicephalus, poseen individuous que
tienen alguna de estas cualidades distintas de callicephalus. La ex-
presion de estas caracteristicas dentro de las poblaciones mas oc-
cidentales de E. brevilineatus es interpretada como evidencia de una
distribucion continua anterior, de poblaciones intermedias la cual
fue interrupida recientemente por alteraciones Pleistocenicas en el
habitat y clima de la parte norte del Desierto Chihuahuense.

A causa de estas observaciones e interpretaciones, la asignacion
taxonomica de los tres especies nominal del Grupo E. brevilineatus
a subespecies de Eumeces tetragrammus (Baird) es recomendada.
Rasgos salientes de coloracion y escamacion de la politipica E. tetra-
grammus claramente la une con Eumeces anthracinus y E. sep-
tentrionalis de el Especie-Grupo E. anthracinus. El Grupo E. an-
thracinus es una de las trece especie-grupos en el genero Eumeces,
y esta asocido aprentemente con un grupo que incluye los escincidos
de Las Especies-Grupos E. fasciatus y E. multivirgatus.

INTRODUCTION

In E.H. Taylor’s (1935b) monograph of the scincid genus
Eumeces, the species were arranged into 1 5 species groups
on the basis of shared color patterns and scalation features.
One of the New World assemblages, the E. brevilineatus
Species Group, included three species: callicephalus Bocourt,

1 879, of Arizona and western Mexico, and brevilineatus Cope,

1 880, and tetragrammus (Baird) 1858, of Texas and northern
Mexico. In contrast to most other species groups erected by
Taylor, species of the E. brevilineatus Group evinced a high

1. Laboratory for Environmental Biology, University of Texas,
El Paso, Texas 79968, USA. Research Associate, Section of Her-
petology, Natural History Museum of Los Angeles County.

Contributions in Science, Number 357, pp. 1-19
Natural History Museum of Los Angeles County, 1985

ISSN 0459-8113

Figure 1. Pattern variation in Eumeces tetragrammus. Top: Young adult (TCWC 45496; Queretaro: El Trapiche); Middle: Old adult with
faded posterior striping (TCWC 36535; Texas: Hidalgo Co.); Bottom: Adult with broad neck lines (TCWC 40751; Coahuila: vie. Cuatro
Cienegas).

degree of intraspecific variation in usually conservative scale
features.

The systematic results of the present study served as a
partial requirement for a Master’s degree at Texas A&M
University (Lieb, 1973), and were partly incorporated into
Conant's (1975) field guide. Taxonomic arrangements ap-
pearing in the latter work thus anticipated the publication of
the present account. Here I document the evidence and ratio-
nale for these proposed taxonomic conclusions and review
the geographic distribution and morphological variation.
Changes are proposed at the intragroup and intergroup levels
within the genus.

MATERIALS AND METHODS

1 have examined over 600 specimens of skinks of the E.
brevilineatus Group. Data concerning individual and geo-
graphic variation were compiled on the following scalation

features (terminology after Taylor, 1935b; Robinson, 1979):
number of postmentals, presence of postnasals, contact of
primary temporals and parietals, enclosure of interparietal
by parietals, number of postlabials, number of longitudinal
dorsal rows, number of latitudinal rows around midbody,
number of nuchal pairs, number of supralabials, number of
supraciliaries, and contact of prefrontals. In addition, data
on individual, geographic, and where possible, ontogenetic
variation in the following color pattern features were also
collected (terminology after Dixon, 1969): expression of me-
dian light line, dorsolateral light lines, dark lateral stripes,
lateral light lines, and upper secondary dark lines. Two as-
pects of gross morphology, axilla-groin/snout-vent length ra-
tio and adpressed limb overlap, were also evaluated. This
data base has been discussed elsewhere (Lieb, 1973), and
only data relevant to the interpretation of relationships among
E. brevilineatus Group taxa are presented here. Moreover,
new data acquired since 1973 have also been incorporated

2 Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus

Figure 2. Pattern of Eumeces brevilineatus and intermediates with E. tetragrammus. Top: Typical short-lined pattern of E. brevihneatus;
Middle: Sympatric zone intermediate form with lengthened body striping (TCWC 39265; Texas: McMullen Co.); Bottom: Possible intermediate
from Sierra Madre foothills (MVZ 185746; Nuevo Leon: vie. Ranchitos).

into this study. Unless otherwise noted, all specimens cited
by museum number have been examined by me.

RELATIONSHIPS BETWEEN
EUMECES BREVILINEATUS AND
E. TETRAGRAMMUS

Eumeces tetragrammus and brevilineatus are distinguished
only by differences in the striped pattern on the body (Taylor,
1 935b). They are much more similar morphologically to each
other than either is to callicephalus. In strongly patterned
tetragrammus, the striping consists of a dark lateral stripe
bordered above and below by light lines (dorsolateral and
lateral light lines). This stripe extends from the neck through-
out the length of the body to the groin or base of the tail (Fig.
1). In brevilineatus, the same striping (lateral stripe, dorso-
lateral, and lateral light lines) terminates on the body just
posterior to the shoulder (Fig. 2). In hatchlings, the dorsum
may be as dark as the lateral stripe. In both tetragrammus

and brevilineatus, the dorsal ground color becomes distinctly
lighter than the lateral stripe with increasing age. In the largest
skinks, the light lines may also fade, resulting in only faint
traces of dorsolateral and lateral light lines. The striping may
also be obscured on the posterior dorsum by prolonged tenure
in formalin; such conversion of fully lined individuals of
tetragrammus to morphs similar to brevilineatus has been
observed in several specimens. Fortunately, even in the most
blackened specimens, enough pattern usually persists to make
positive identification possible.

Taylor (1935b) found no evidence of intergradation be-
tween tetragrammus and brevilineatus, but his distributional
data suggested a substantial zone of geographic sympatry in
Texas. Later (Taylor, 1943) he reported a specimen of bre-
vilineatus from Tamaulipas, Mexico, thus indicating an ex-
panded zone of overlap with tetragrammus that included not
only southern and central Texas (from Burnet Co. south-
ward), but also a large section of northeastern Mexico. This
broad zone of overlap was further substantiated by Smith

Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus 3

Figure 3. Hatchling Eumeces tetragrammus (TCWC 41555-61) from a clutch found with a fully lined female, 2 mi. W Bruni, Webb Co.,
Texas (TCWC 3927 1 ). Note the dark coloration and the incomplete expression of the dorsolateral light lines. Photograph by Richard J. Baldauf.

( 1 946), who provided distribution maps that indicated over-
lap; Brown ( 1 950), who reported a specimen of tetragrammus
from Bexar Co., Texas (immediately south of the Edwards
Plateau), and specimens of brevilineatus from Cameron and
Hidalgo cos. (in the lower Rio Grande Valley); Conant (1958),
who published distribution maps indicating the overlapping
ranges; Anderson (1962), who reported a specimen of bre-
vilineatus from San Luis Potosi; Holman (1968), who re-
ported a Pleistocene fossil tetragrammus from Kendall Co.
(on the Edwards Plateau); Raun and Gehlbach (1972), who
published a county distribution map for brevilineatus in Tex-
as with a record for Cameron Co. (probably that of Brown,
1 950), and examined specimens from Hidalgo Co. Raun and
Gehlbach (1972) also accepted the Bexar Co. tetragrammus
record, rejected literature records for western and northern
Texas, and doubted the validity of the central Edwards Pla-
teau Burnet Co. record.

Strecker (!909a) reported tetragrammus from an isolated
locality in Burnet Co., to the north of the range. Taylor ( 1 935b)
questioned this record, apparently a single specimen in the
Strecker Museum. The specimen is not now in the museum

nor are there records that it was preserved, retained or ex-
changed (pers. comm., D. Lintz, Strecker Museum). It could
have been a misidentified septentrionalis. The two forms are
superficially similar, and it is clear elsewhere that Strecker
( 1 908, 1 909b; Strecker and Williams, 1 927) could not readily
distinguish the two.

Taylor (1943) reported a very small specimen of brevilin-
eatus from the coastal plain of Tamaulipas. This individual
(UIMNH 22443) was found to be a desiccated hatchling of
tetragrammus in which the light lines are poorly defined and
partially obscured by skin creases. Hatchlings often have such
truncated light lines (Fig. 3).

Brown ( 1 950) did not cite specimen numbers or museums
for Texas records, and the reports of tetragrammus from
Bexar Co. and E. brevilineatus from Hidalgo and Cameron
cos. are unverifiable. I have not seen specimens of tetra-
grammus from Bexar Co., but I have examined a series of
brevilineatus reportedly from Cameron Co. (FMNH 2721 5—
17). Brown could have utilized these specimens, as he did
examine some FMNH holdings. These, however, are the only
representatives of the short-lined form from the lower Rio

4 Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus

Grande Valley seen by me. The specimens examined in the
B.C. Brown private collection from Hidalgo Co. that were
catalogued as brevilineatus (BCB 2402, 2 specimens) are aged
tetragrammus with faint light lines. Similarly, Anderson's
(1962) record of brevilineatus for San Luis Potosi (AMNH
66999) is a specimen of tetragrammus that has been over-
preserved in formalin.

The allocation of a Pleistocene fossil to tetragrammus was
based upon dentary tooth characters (Holman, 1968). Al-
though I have not examined the fossil material (Univ. Texas
Bureau Econ. Geol. 40450-1666), the allocation is dubious
in that study of several cleared and stained specimens of both
taxa does not corroborate Holman’s observed differences in
the spacing and robustness of the teeth. Furthermore, even
if tetragrammus occurred in the area during the Pleistocene,
it does not now occur there.

The county-based maps and distributional summaries of
Raun and Gehlbach (1972) were based upon literature rec-
ords, examined specimens, and in some cases, museum cat-
alogue files. The presumed distributional errors for brevili-
neatus of Brown (1950) were thus perpetuated, and other
museum records from the lower Rio Grande Valley of Texas
were also accepted. Besides the BCB specimens from Hidalgo
Co. mentioned above, a misidentified series from this county
at a second museum (TCWC 18169-73, 18176-82, verified
as tetragrammus by me) was also apparently utilized by Raun
and Gehlbach (1972) for the brevilineatus distribution map.

Thus, extant material with acceptable locality data indicate
that the major area of sympatry of brevilineatus and tetra-
grammus occurs in the coastal plain of southern Texas, north
of the Rio Grande Valley and south of the Edwards Plateau
escarpment. Plotting of locality data (Fig. 4) further suggests
that actual sympatry in southern Texas occurs only in the
vicinity of the Nueces Riverdrainage system, from the mouth
of the river west and then north to the edge of the Pleateau.
The two forms may also occur together in the Rio Grande
drainage in the vicinity of Laredo (Webb Co.). There is a
specimen of brevilineatus from this area (UMMZ 1 14253),
and Werler (1951) reported on the hatchlings of a clutch of
eggs from a female tetragrammus from Laredo (specimens
not examined). Nevertheless, these regions mark the north-
ernmost limit of the range of tetragrammus. and the southern
extent of the Gulf Coastal Plain populations of brevilineatus.
E. brevilineatus do occur in northern Mexico, but are known
only from the Sierra del Nido of Chihuahua and the Coahuila
Folded Belt of Coahuila and Nuevo Leon. These populations
occur in rocky habitats within pinyon-juniper, oak woodland,
or piedmont areas similar to those in western Texas. Within
the northern Chihuahuan Desert, brevilineatus may occur in
low desert mountain ranges where the appropriate mesic hab-
itats are found. On the other hand, tetragrammus are pri-
marily associated with the coastal plain lowlands, particu-
larly riparian or mixed grassland-brushy areas with sandy
substrata. Toward the southern part of its range in Mexico,
the species also occurs in rocky habitats in the foothills of
the Sierra Madre Oriental and in the isolated coastal plain
ranges of Tamaulipas (Sierra de Tamaulipas, Sierra San Car-
los). The distributional ranges of tetragrammus and brevilin-

eatus should make contact in northeastern Nuevo Leon, in
areas where the foothill habitats of the Coahuila Folded Belt
and northern Sierra Madre abut those of the Tamaulipan
Coastal Plain. However, only two specimens are known from
this region. Their significance is discussed below.

A presumably disjunct population of tetragrammus occurs
in the Cuatro Cienegas Basin of Coahuila (Zweifel, 1958);
the four specimens from this population (AMNH 77316,
TCWC 40750-52) are all from riparian or other mesic hab-
itats in the Moor of the basin. The two adult specimens are
distinctive in having comparatively wide dorsolateral light
lines on the neck (Fig. 1 ), and a dorsal ground color of a pale
shade of gray or gray-brown. The other two specimens are
juveniles with coloration and color pattern similar to other
tetragrammus populations; the scalation of all four individ-
uals is fairly typical of the species (Table 1). E. brevilineatus
is known to occur in a low mountain range approximately
75 km (air) north of the Cuatro Cienegas area, but because
of the habitat specificity described above, such desert range
populations are almost certainly isolated from the riparian
enclave of tetragrammus in the Cuatro Cienegas Basin. The
occurrence of a disjunct population of tetragrammus in this
basin is thus interesting in a biogeographic sense, but does
not appear to represent sympatry with brevilineatus. Other
taxa of obvious Tamaulipan affinities are also found in the
Cuatro Cienegas area (e.g., the snakes Drymarchon corais
and Drymobius margaritiferus ), as are riparian species with
close relatives in the Gulf Coast lowland/Salado River drain-
age to the east (see Morafka, 1977).

What would otherwise be a reasonably clear-cut habitat
difference between the two species breaks down south of the
Edwards Plateau in south-central Texas. In this area, and
southward into the zone of sympatry with tetragrammus.
brevilineatus occupy non-rocky brushland, grassland, and ri-
parian habitats that are essentially identical to those utilized
by tetragrammus. Moreover, within this area of sympatry,
color pattern intermediates have been observed. In the in-
termediates, the body striping pattern extends posterior to
transverse dorsal row 30 and terminates at or before the
midbody region (Fig. 2). In brevilineatus outside the sym-
patric zone, such elongated stripes do not occur. In tetra-
grammus. both within and outside of the sympatric zone,
the body striping is continuous from neck to groin, although
it may be somewhat faded posterior to the midbody in old
adults (Fig. 1). As mentioned earlier, some tetragrammus
hatchlings have shortened body lines. In these individuals,
however, the fully lined pattern seems to develop ontoge-
netically before a snout-vent length of 45 mm is reached.
Even though most small juveniles of tetragrammus are fully
lined, the allocation of individual skinks to intermediate sta-
tus is thus possible only for adults and subadults over 45
mm in snout-vent length.

In a series of six specimens from southern Live Oak Co.
(TAIC), one is fully lined (tetragrammus. No. 1 1 7), four are
“short” lined (brevilineatus. 258. 1 -.2, 123.2-.3), and the sixth
is intermediate (123.1). A second sample, a series of five
individuals from the northwestern edge of the sympatric zone
in south-central Uvalde Co. (TCWC), contains one tetra-

Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus 5

Figure 4. Distribution of Eumeces brevilineatus Group taxa in southwestern North America. Circles indicate Eumeces tetragrammus ; squares
E. brevilineatus; triangles E. callicephalus. Stippled areas mark the regions of presumed sympatry between brevilineatus and tetragrammus,
the circle within squares indicating localities where both forms are known to occur together, and arrows indicating populations that contain
intermediate forms (see text).

grammus (44175), two brevilineatus (44173-74), and two
intermediates (44171-72). The fully lined individual (a fe-
male) was taken under a rock in copulo with one of the short-
lined males (R.A. Thomas, pers. comm.). The two lizards
obligingly repeated their act in the laboratory for photographs
(Fig. 5), and the eight subsequent eggs produced only fully
lined offspring (TCWC 44176-83).

Other intermediates between brevilineatus and tetragram-
mus are known from individual specimens collected in Jim
Wells Co. (KU 88 1 2), LaSalle Co. (TAIC 643), Live Oak Co.
(TCWC 10537), and McMullen Co. (LACM 134855, TCWC
39265, TNHM 28836); their occurrence spans most of the
coastal plain drainage of the Nueces River. In addition to
the typical short-lined specimens of brevilineatus from Live
Oak (TAIC) and Uvalde (TCWC) cos., other short-lined in-
dividuals within the Nueces drainage have been examined
from Dimmit Co. (KU 8195), McMullen Co. (TCWC 39266-

67), Nueces Co. (TCWC 18175), and elsewhere in Live Oak
Co. (TCWC 10535-36, 10538). A fully lined tetragrammus
is known from Frio Co. (CM 10558). Over the entire Nueces
River drainage area, the total observed numbers of fully lined,
intermediate, and short-lined individuals were 3, 9, and 13,
respectively. Although the short-lined form seems to pre-
dominate, there are still relatively few specimens available
from this poorly sampled region.

As mentioned previously, another area of potential contact
of tetragrammus and brevilineatus is in northeastern Nuevo
Leon. Only two specimens (MVZ) are known from this re-
gion, both from the same locality about 35 km SE of Ciudad
Monterrey. One specimen (185745) is a typical fully lined
tetragrammus, the other (185746) has the shortened light
lines that are typical of the condition in south Texas inter-
mediate populations (Fig. 2). In this “intermediate” speci-
men, however, there are distinct traces of a complete dark

6 Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus

Table 1. Geographic variation in six characters for the nominal forms of Eumeces tetragrammus. Frequencies are followed (in parentheses)
by sample sizes. Letters correspond to geographic regions mapped in Figure 6. Asterisks indicate a condition that is expressed on at least one
side of the head.

Postmental

divided

Postnasals

present*

Interparietal

enclosed

Primary

temporal

contacts

parietal*

Postlabials

single*

Nuchal Y-mark
present

A: NE Mexico
tetragrammus

0.10(51)

0.04 (51)

0 (51)

0.18 (50)

0.03 (39)

0 (42)

B: Cuatro Cienegas

tetragrammus

0 (4)

0 (4)

0 (4)

0.25 (4)

0.25 (4)

0 (4)

C: South Texas

tetragrammus

0.15 (74)

0.01 (74)

0 (73)

0.19 (72)

0.06 (71)

0 (68)

D: Sympatric zone

0.03 (33)

0.10 (32)

0 (24)

0.25 (24)

0.03(31)

0 (13)

E: West-central Texas

brevilineatus

0.05 (151)

0.07 (149)

0.02 (149)

0.21 (149)

0.14(139)

0.01 (68)

F: East-central Texas

brevilineatus

0.18 (33)

0.21 (33)

0.03 (33)

0.30(33)

0.26 (35)

0 (23)

G: Coahuila-West Texas
brevilineatus

0.17 (47)

0.09 (45)

0.04 (47)

0.31 (45)

0 (47)

0.60 (47)

H: Sierra del Nido
brevilineatus

0 (4)

0 (4)

0.33(3)

0.67 (3)

0.33(3)

0.25 (4)

I: USA-NW Mexico

callicephalus

0.96 (80)

0.58 (79)

0.64 (80)

0.67 (76)

0.99 (67)

0.87 (61)

J: West Mexico

callicephalus

0.59 (41)

0.83 (41)

0.89 (38)

0.87 (39)

0.96 (28)

0.77 (30)

lateral stripe extending posterior to the termination of the
light lines. Such extension of the dark lateral band was not
observed in any of the southern Texas intermediates, al-
though such a characteristic could be obscured in some in-
dividuals by a long period of preservation. Nevertheless, in
the absence of more information on the distribution of short-
lined forms in the region, the assignment of the specimen to
intermediate status is provisional.

Evidence for conspecificity of the forms tetragrammus and
brevilineatus arises from the following observations: 1) the
extremely close morphological similarity between the two
forms in southern Texas; 2) the apparent lack of behavioral
pre-mating isolating mechanisms where the two occur in
sympatry; and 3) the presence of color pattern intermediates
in the sympatric zone. This evidence is further supported by
the lack of significant overlap in the distributions of the two
forms, and by a lack of ecological segregation in the principal
area of geographic contact and sympatry.

Alternatively, the two forms could be acting as parapatric
species that only rarely hybridize. As noted before, copula-
tion of a short-lined male and a long-lined female were ob-
served under field and laboratory conditions, and that the
resultant offspring were all fully lined. Assuming that the
short-lined individual was indeed the male parent, then the
expression of the long-lined pattern of tetragrammus would
seem to be dominant over the short-lined pattern of brevi-

lineatus. Such a dominance relationship in the inheritance
of color pattern, however, does not readily explain how the
intermediate condition arises, or why there is an apparent
preponderance of short-lined individuals in the sympatric
zone. On the other hand, should the gene pools of brevilin-
eatus and tetragrammus be separated by post-mating isolat-
ing mechanisms, then the presence of occasional interme-
diates in the contact zone might represent sterile F,’s incapable
of backcrossing to the parental stocks.

My studies on brevilineatus and tetragrammus clarify some
aspects of the relationships of the two forms (particularly in
distribution), but offer little in the way of an objective de-
cision as to their specific status. From the material I have
examined and the characters 1 have inspected, I feel that for
the present the two nominal forms should be considered
subspecies of a single species. Full resolution of their status,
however, will require larger samples from the contact zone
and a complete genetic analysis of the populations involved.

DISTRIBUTION AND STATUS OF
EUMECES CALUCEPHALUS BOCOURT

When compared with both tetragrammus and brevilineatus,
callicephalus differs significantly in several aspects of cephalic
scutellation. These characters are as follows (frequencies of
tetragrammus and brevilineatus in parentheses, respectively;
N > 100 in all cases): postmental scale divided, 0.91 (0.14

Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus 7

Figure 5. Male brevilineatus (TCWC 44173) and female tetragrammus (TCWC 44175) copulating under laboratory conditions. Both indi-
viduals are from 8 mi. N Uvalde, Uvalde Co., Texas; they were also observed copulating when captured. Photograph by Robert A. Thomas.

Figure 6. Geographic areas for combined samples indicated in Ta-
ble 1. A, B, C: Eumeces tetragrammus; D: principal sympatric zone
of E. tetragrammus and E. brevilineatus in southern Texas; E, F, G,
H; /.'. brevilineatus; I, J: E. callicephalus.

and 0.08); postnasal scales present on at least one side of the
head, 0.68 (0.03 and 0.09); enclosure of interparietal scale
by parietals or azygous scales, 0.62 (0 and 0.03); primary
temporal contacting parietal on at least one side of the head,
0.77 (0. 1 7 and 0.25); and single (rather than double or triple)
postlabials on at least one side, 0.97 (0.05 and 0.12). How-
ever, the frequencies of these characters vary geographically
(Table 1, Fig. 6) and are discussed further below.

Most of the vividly patterned specimens of callicephalus
vary from the basic tetragrammus/brevilineatus stripe pat-
tern in having a persistent dark lateral stripe from neck to
groin, faded dorsolateral and lateral light lines posterior to
the shoulder, and a light median line on the anterior dorsum
that bifurcates on the nuchal scales (Fig. 7). In some adults,
particularly those from the southern part of the range, the
color pattern is faded, with concomitant loss of the bifur-
cating Y-mark and median light line, and a general obscuring
of the lateral stripe and other light lines. Such faded speci-
mens seem to have been the basis for the original recognition
of humilis Boulenger 1887, a name based on patternless in-
dividuals of callicephalus (Robinson, 1979).

8 Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus

Figure 7. Pattern of Eumeces callicephalus and western E. brevilineatus. Top: E. callicephalus with typical pattern, including nuchal Y-mark
(BYU 14260; Chihuahua: Cuiteco); Middle: E. brevilineatus from western part of range, with faint nuchal Y-mark present (UAZ 16816;
Coahuila: vie. Piedra Blanca); Bottom: E. callicephalus with faded dorsal pattern (UTEP 4865; Sinaloa: vie. Mazatlan).

The geographic range of callicephalus in Mexico has been
somewhat overestimated. Bocourt (1879) described the
species from a specimen (MNHP 1643, not examined) sent
to him in the mid-1800’s by Alfredo Duges of Guanajuato,
Mexico. Later authors, particularly Cope (1887) and H.M.
Smith and Taylor (1945), assumed the city of residence of
the collector to be the type locality of the species. There are
also two specimens of callicephalus studied by Cope (ANSP
1 3604-05) that bear tags reading only “Guanajuato, Mexico
. . . Duges.” The original type description, however, does not
clearly give the collection locality as the city or state of Gua-
najuato, but only associates the specimen with the collector
who resided there. Additional information is contained in a
paper by Duges (1889) on a comparison of the herpetofauna
of the Guanajuato region with that of the Guadalajara (Ja-
lisco) area. Here, he specifically associates callicephalus with
the Jalisco capital, citing lynxe as characteristic of his own
locale. This condition still pertains, and it is likely that the
type specimen of callicephalus (as well as the ANSP material)
originated from near Guadalajara or from some other locality

farther to the west. Taylor (1935b) indicated that the re-
maining material in the Alfredo Duges Museum in Guana-
juato bears the label “San Bias,” presumably referring to the
coastal town in Nayarit. Specimens of callicephalus are known
from within 50 km of Guadalajara.

E. callicephalus has also been reported from Queretaro,
Mexico (Smith and Taylor, 1945). H.M. Smith (Univ. Col-
orado, pers. comm.) has advised me that this is another of
Duges’s specimens, but I have been unable to locate either
the original citation or the specimen. The locality is “Huax-
teca Potosina,” which is more applicable to a general region
in the northeastern part of the state than to any specific
locality. E. tetragrammus occurs in the Huaxteca area, and
the report could be based upon a specimen of this species.

Two additional localities, records of Taylor ( 1935b), have
already been questioned and corrected: Tombstone, Arizona
(see Zweifel, 1962) and "Ciudad” (=La Ciudad de Rocas) in
Durango. The latter record, based upon a specimen (British
Mus. Nat. Hist. 83.413, not examined) collected by Forrer
in the early 1800's, is a callicephalus (in litt., R.G. Webb,

Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus 9

UTEP). However, La Ciudad is higher in elevation (more
than 2400 m) than other records for the species in western
Mexico, and only brevirostris occur there. Conant (1969:86)
has discussed problems involving confused localities for oth-
er specimens credited to Forrer, and it seems likely that this
specimen actually came from a much lower elevation some-
where along the trail between Ventanas and La Ciudad. Thus,
the geographic range of cal/icepha/us is confined to low and
moderate elevations west of the Continental Divide, extend-
ing from the general area west of Guadalajara north to south-
ern Arizona and adjacent New Mexico.

The geographic range of callicephalus is allopatric to those
of the other Eumeces brevilineatus Group members. E. bre-
vilineatus occur in the Sierra del Nido of Chihuahua, ap-
proximately 140 km east of the nearest locality for cal-
licephalus (Chihuahua: 8 mi. W Matachic, AMNH 68295).
This Chihuahua brevilineatus population is known from four
specimens (MVZ 70702-03, LACM 116401, UTEP 62), all
of which were collected in Canon de Santa Clara on the
eastern slope of the range. The LACM and MVZ specimens
have the typical color pattern of brevilineatus, whereas the
UTEP specimen has traces of the ca/licephalus-like bifur-
cating head lines on an otherwise typical brevilineatus pat-
tern. Such traces also occur in some individuals of brevilin-
eatus from the western part of the range, but do not occur
in eastern populations nor in tetragrammus. One specimen
(MVZ 70702) is badly damaged, and most scalation features
could not be determined (postmental entire, postnasals ab-
sent). In MVZ 70703, the postmental is entire, the postnasals
absent, the interparietal not enclosed, the primary temporal
contacts the parietal on the right side, and the postlabials are
double. The LACM specimen is similar, except that neither
primary temporal contacts a parietal. UTEP 62 has an entire
postmental, no postnasals, an enclosed interparietal, contact
of the primary temporal with the parietal on the left side,
and a single postlabial on the right side.

The small series from the Sierra del Nido thus contains
two individuals (MVZ 70702, LACM) that are typical of
eastern brevilineatus populations in color pattern and scal-
ation, one (UTEP) that contains a mixture of brevilineatus
and callicephalus color pattern and scalation similarly con-
tained in western brevilineatus, and one (MVZ 70703) that
has the typical brevilineatus color pattern and approaches the
callicephalus scalation condition only in the contact of a
primary temporal and a parietal (occurs at an overall fre-
quency of 0.77 in callicephalus and 0.25 in more easterly
brevilineatus). A similar trend is seen in other western bre-
vilineatus populations (ca. 250-450 km to the east) for which
larger series are available. Overall in the samples from Trans-
Pecos Texas and Coahuila (Fig. 6; Table 1), the Y-shaped
bifurcating head lines occur in 60 percent (N = 47) of the
individuals, and the frequencies of two of the five scale char-
acters typical of callicephalus are higher than those in the
central Texas brevilineatus populations. Slightly higher fre-
quencies of ca/licephalus-hke scale features also occur in bre-
vilineatus populations in the northeastern Edwards Plateau
and along its southeastern periphery (Fig. 6, Table 1). The
bifurcating head lines observed in many western brevilinea-

tus, however, do not occur in the more easterly populations,
nor in tetragrammus.

These data indicate a pronounced tendency for the west-
ernmost populations of brevilineatus to express a color pat-
tern feature otherwise more characteristic of callicephalus,
and suggest similarities in some scalation features as well.
The Trans-Pecos and Coahuila populations of brevilineatus,
however, are separated from the range of callicephalus by
400 km of unsuitable Chihuahuan Desert habitats. There are
no authenticated distributional data for callicephalus east of
the Continental Divide, so it is unlikely that the observed
trends in western brevilineatus populations could be due to
hybridization with the other form. These patterns do suggest,
however, that intermediate populations between brevilinea-
tus and callicephalus probably extended across much of the
now intervening desert during the pluvial past. The estimated
extent of pinyon-juniper woodlands during the Wisconsin
glacial maximum (ca. 20,000 years B.P.) is to elevations as
low as 300-600 m at the 25th parallel, and 600-900 m at
the 30th parallel (see Morafka, 1977, for review). Such dis-
placements would have provided suitable corridors for these
skinks across much of Chihuahua and Coahuila north of the
Arteaga and Parras anticlines. Thus the differentiation of
callicephalus and brevilineatus may be the result of interrup-
tion of a continuous gene pool by climate and vegetation
changes of the last 10,000 years. Further studies on the vari-
ation and distribution of both brevilineatus and callicephalus
in Chihuahua are needed, particularly in regard to the dis-
continuous areas of pinyon-juniper and oak habitats from
which neither form is known. The degree of similarity of
western Texas and Coahuila populations of brevilineatus to
callicephalus, however, suggests that the two forms should
be considered allopatric members of a single species, and
that callicephalus be retained as a distinct subspecies.

TAXONOMIC AND DISTRIBUTIONAL SUMMARY

The three forms recognized as distinct species of the Eumeces
brevilineatus Group of Taylor ( 1 935b) are herein considered
subspecies that are distributed in a discontinuous arc from
western Mexico north to the southwestern United States and
south again into eastern Mexico. The oldest available name
for the three taxa is Eumeces tetragrammus (Baird). The
following accounts summarize information on variation and
distribution for the species and its three subspecies.

Eumeces tetragrammus (Baird)

Plestiodon tetragrammus Baird, 1858:256.

Eumeces callicephalus Bocourt, 1879:431-433.

Eumeces brevilineatus Cope, 1880:18-19, 44, 46.

Eumeces humilis Boulenger, 1887:377.

[Eumeces tetragrammus ] var. funebrosus Cope, 1900:630,

661.

Type specimens. The type series originally consisted of 12
or more specimens from Matamoros, Tamaulipas (USNM
3124; Taylor, 1935b). Taylor designated 3124A as the lec-

10 Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus

totype (now USNM 165662, examined; the remaining spec-
imens in the original type series were not seen).

Diagnosis. Maximum snout-vent length 74 mm, dorsal
scales around body subequal, in 26 or 28 parallel rows at
midbody; scale lying medial to postgenial scale longer than
wide (see Robinson, 1979); supraoculars four; body striping
present in all but old adults, striping terminates anterior to
the midbody region or on tail within three to five scales
posterior to vent; single dark lateral stripe at least two scale
rows wide present on each side, bordered above and below
by light lines; dorsolateral light lines occupy third and fourth,
or fourth only, lateral scale rows of neck; lateral light line
passes through auricular opening; median light line, if pres-
ent, bifurcates on the nuchal scales and extends posteriorly
no more than a third of the body length; distal portions of
tails in hatchlings and juveniles bright blue.

Variation. Supranasal scales usually in contact; prefrontals
in contact or not; parietals may or may not enclose inter-
parietal; postnasals present or absent; supraciliaries vary from
six to nine, usually seven or eight; postsuboculars vary from
two to four, usually three; postlabials single or double; post-
mental single, or divided by a transverse suture; dorsal scales
from occiput to above vent 52-60; lamellae under fourth toe
of hind limb 10-18; upper secondary dark lines (see Dixon,
1969) present or absent; complete pattern loss may occur in
very large or old adults.

Most patterned individuals may be identified to subspecies
as follows:

la. Dark lateral stripe extends from axilla to groin ... 2
1 b. Dark lateral stripe terminates anterior to midbody region

E. t. brevilineatus

2a. Postlabials single; median light line anteriorly bifurcat-
ing on nuchals to form a Y-shaped head marking . . . .

E. t. callicephalus

2b. Postlabials double; median line absent, no bifurcating

lines on nuchals E. t. tetragrammus

Distribution. Southwestern United States, northeastern and
western Mexico (Fig. 4).

Eumeces tetragrammus tetragrammus (Baird)

Type specimens. See species account.

Diagnosis. Postlabial scales usually double; postnasals usu-
ally absent; interparietal usually not enclosed by parietals;
dark lateral stripes, dorsolateral and lateral light lines present
throughout body length in adults and juveniles; median light
line absent.

Variation. Some hatchlings have poorly expressed or trun-
cated light lines. With the exception of the population at
Cuatro Cienegas, Coahuila, there is little geographic variation
in color pattern. As noted earlier, adults from this area have
a relatively pale dorsal color and slightly wider dorsolateral
light lines on the neck (Fig. 1). These characteristics are not
evident, however, in the two juveniles from the same area.
Additional specimens are needed to evaluate the ontogeny
of color pattern in this unique geographic variant.

Distribution. See Figure 4.

Habitat. This subspecies is most abundant in brushlands
and grasslands with sandy substrata, but it also occurs in
tropical deciduous forest, palm forest, subtropical brushlands
with rocky substrates, and in mesic forests associated with
riparian areas. The known elevational range is sea level to
1060 m.

Eumeces tetragrammus brevilineatus (Cope)

Type specimens. There are two syntypes from Helotes,
Bexar Co., Texas (USNM 101 59A and 101 59B). Taylor
( 1 935b) designated 101 59B as the lectotype.

Diagnosis. Postlabials usually double; interparietal usually
not enclosed by parietals; body striping terminates between
shoulder and midbody.

Variation. Ground color in adults varies from dark gray
to greenish-gray to brown, with or without dark edges on the
dorsal scales. Specimens from far western populations often
possess a light bifurcating mark on the nuchals (Fig. 7).

Distribution. See Figure 4.

Habitat. Eumeces t. brevilineatus is most abundant in xe-
rophilous woodlands with rocky substrata. It also inhabits
grasslands and brushlands with sandy substrata and riparian
woodlands through xeric areas. Reported elevations range
from about 150 m to 2300 m.

Eumeces tetragrammus callicephalus (Bocourt)

Type specimen. The holotype (MNHP 1643, not ex-
amined) was sent from Guanajuato to Paris by Alfredo Duges
in 1868, but it was probably collected elsewhere.

Diagnosis. Postlabials usually double, postnasals usually
present, dark lateral stripe usually present throughout body
length, light bifurcating head lines and short median light
line usually present.

Variation. Color of adults in preservative is gray or gray-
green dorsally, with dark brown to red-brown lateral stripes.
The median light line, and the dorsolateral and lateral light
lines, may be faded or absent (Fig. 7).

Distribution. See Figure 4.

Habitat. This subspecies seems to be most abundant in
wooded rocky canyons in the northern part of its range, usu-
ally within an elevational range of 900 to 1700 m. In south-
western Mexico, they occur in mesic foothill forests and trop-
ical deciduous lowland habitats. In the southern part of the
range it is replaced at higher elevations by Eumeces brevi-
rostris and by E. multilineatus in the north. Habitats on the
eastern slopes of the Sierra Madre Occidental in Durango
and Zacatecas, as well as across the southern margin of the
Mexican Plateau that would appear suitable for E. t. calli-
cephalus, are inhabited by E. lynxe. Suitable habitats in west-
ern New Mexico and western Chihuahua are apparently un-
occupied.

SPECIES GROUP STATUS

Taylor (1935b) defined 15 species groups of Eumeces, but
did not fully indicate his criteria for recognizing all groups.
Moreover, the relationships between the species groups were

Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus 1 1

largely unstated, and were essentially confined to a phylo-
genetic tree (p. 38) with little comment or explanation. It is
clear from this tree, as well as from the key to the species
and from the text, that Taylor’s groups were based upon
shared features of color pattern and/or scalation. How he
arrived at many of his conclusions is not easily determined.

Taylor’s phylogeny indicated that the E. brevilineatus
Group (i.e., Eumeces tetragrammus as defined above) was
closely related to the E. fasciatus Group, an assemblage of
12 nominal species widely distributed in eastern Asia and
the southeastern United States. The unifying feature of the
E. fasciatus Group is primarily the presence of five dorsal
light lines on the body and tail in all juveniles and many
adults. In tetragrammus, only parts of this E. fasciatus- type
pattern are present: the median light line is absent (or present
only anteriorly), and the paired light lines are absent on the
distal tail and/or reduced on the body. E. fasciatus Group
species also differ from tetragrammus in the following (char-
acteristics of the latter in parentheses): larger body sizes at-
tained, with snout-vent lengths in excess of 80 mm typical
of most species (observed snout- vent maximum 76 mm);
interspecific variation in scales around body ranging from 22
to 26 (26-28 intraspecifically); postnasals usually present in
eight of the 12 species (usually present in one of three sub-
species). However, in these features of color pattern, body
size, and scalation, tetragrammus is much less similar to any
E. fasciatus Group form than it is to members of the E.
multivirgatus and E. anthracinus species groups.

The E. multivirgatus Group was defined by Taylor (1935b)
as follows: multivirgatus, gaigei, humilis, parviauriculatus,
and parvulus. Subsequent studies have considerably altered
the original composition of the species group. E. gaigei is
now a junior synonym of multivirgatus, and the gaigei spec-
imens that Taylor utilized are now E. m. epipleurotus (Tay-
lor, 1935a; Axtell, 1961; Mecham, 1980). E. humilis was
based on patternless multivirgatus from the United States
(H.M. Smith, 1942; Mecham, 1957), and on patternless cat-
licephalus from Mexico (Robinson, 1979). Robinson (1979)
also removed parviauriculatus and parvulus from the E. mul-
tivirgatus Group and allocated them to the E. brevirostris
Group (sensu Dixon, 1969). One species, multilineatus (known
only from a small area in the high elevations of the Sierra
Madre of Chihuahua), was added to the E. multivirgatus
Group at the time of its description (Tanner, 1957). It is
similar to multivirgatus, and was independently described as
E. multivirgatus mexicanus Anderson and Wilhoft, 1959 (see
Legler and Webb, 1960). The present composition of the E.
multivirgatus Group consists of only multilineatus and mul-
tivirgatus.

Individuals of multilineatus have the fully developed five-
lined dorsal pattern characteristic of the E. fasciatus Group,
as do juveniles of some populations of the polytypic species,
multivirgatus. Most of the populations of the latter form,
however, have modified five-lined patterns in adults. The
predominate modifications are patternless (unicolor), four-
lined, and five-lined patterns that lack the nuchal Y-mark,
as well as the multiple-lined pattern, the source of the sci-
entific and common names (Many-lined Skink). E. tetra-

grammus resembles the species of the E. multivirgatus Group
in body size and scalation features, and the color pattern of
E. t. tetragrammus and the four-lined multivirgatus are sim-
ilar. However, the great variation in color pattern in the E.
multivirgatus Group and the tendency for reduced number
of scales around the body (24) are absent in tetragrammus.

Greater resemblance to tetragrammus is found with the
E. anthracinus Group. As defined by Taylor, the E. anthra-
cinus Group contained three species: anthracinus. copei, and
septentrionalis. At present, however, only septentrionalis and
anthracinus remain in this group; copei was removed by
Dixon (1969) and placed in the E. brevirostris Group. E.
septentrionalis is a grassland species that is locally abundant
over much of central North America from Manitoba to coast-
al Texas; anthracinus is a relatively rare forest species dis-
tributed discontinuously in the eastern United States. The
principal diagnostic characters for the two (P.W. Smith and
H.M. Smith, 1952) are the placement of the light lateral line
in the auricular region (through the ear opening in anthra-
cinus, above it in septentrionalis) and the condition of the
postmental scale (entire in anthracinus, divided in septen-
trionalis). E. tetragrammus resembles these species in their
essentially four-lined color patterns, comparable body size
and proportions, and similar scalation (including 26-28 scale
rows around body). Moreover, the position of the body strip-
ing is identical in anthracinus, septentrionalis, and tetra-
grammus, except the striping extends well onto the tail in-
stead of terminating at the shoulder (E. t. brevilineatus) or
within two to five scales posterior to the vent ( E . t. tetra-
grammus, E. t. callicephalus). Some individuals of both E.
anthracinus Group species possess traces of middorsal light
lines as well. When present, this line occurs as an indistinct
postnuchal light stripe with or without a dark border; no
trace of a bifurcating Y-mark is ever present.

The resemblance of E. tetragrammus to the E. anthracinus
Group is so marked that continued placement of tetragram-
mus in a separate species group is unwarranted. The three
species, anthracinus, septentrionalis, and tetragrammus rep-
resent a fairly homogenous group within the genus, partic-
ularly when compared with such diverse assemblages as the
E. fasciatus, E. multivirgatus, and E. brevirostris groups.

In the nearly five decades since the appearance of Taylor’s
generic monograph, a substantial number of taxonomic
changes have occurred in the species or species groups of
Eumeces. Only one of the subsequently introduced species
names ( multilineatus ) has persisted as part of the North
American fauna. Most studies have resulted in the reallo-
cation of various taxa to other species or species groups. In
order to summarize the relationships of tetragrammus, I pro-
vide in Table 2 a revised group classification of the genus
Eumeces. This classification and species group terminology
is in large based upon the arrangement of Taylor (1935b:35-
39), but it incorporates subsequent changes. Additionally, I
have incorporated the following new changes: the use of
subgeneric nomenclatural categories for species series and
sections above the level of species groups; the E. longirostris
Group and the E. obsoletus Group are combined into the
same species series. Taylor placed the E. longirostris Group

12 Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus

Table 2. A classification of the genus Eumeces (modified from Tay-
lor, 1935b). Pertinent literature is indicated by the citations in pa-
rentheses.

Genus Eumeces
Eumeces Section
E. schneiderii Species Series

E. schneiderii Species Group (Eiselt, 1940; Mertens, 1946)*

E. algeriensis
E. pavimentatus
E. princeps
E. schneiderii

E. taeniolatus Species Series
E. taeniolatus Species Group
E. poonaensis (Sharma, 1970)

E. taeniolatus

E. schwartzei Species Group
E. altamirani
E. managuae
E. schwartzei
Pariocela Section
E. obsoletus Species Series
E. longirostris Species Group
E. longirostris
E. obsoletus Species Group
E. chinensis

E. coreensis (Doi and Kamita, 1937; Smith et al., 1975)

E. kishinouyei
E. obsoletus( Hall, 1976)

E. lynxe Species Series

E. lynxe Species Group (Parker, 1960; Webb, 1968)

E. lynxe

E. sumichrasti Species Group (Smith and Etheridge, 1953)

E. sumichrasti
E. fasciatus Species Series
E. anthracinus Species Group
E. anthracinus (Smith and Smith, 1952)

E. septentrionalis (Smith and Slater, 1949)

E. letragrammus

E. fasciatus Species Group (Davis, 1969; Hikada, 1978a; Mur-
phy et al., 1983)

E. barbouri

E. capito (Smith et al., 1975)

E. elegans

E. fasciatus

E. inexpectatus

E. laticeps

E. latiscutatus

E. marginatus

E. okadae (Hikada, 1978b)

E. oshimensis
E. stimsoni

E. tamdaoensis (Bourret, 1937)

E. tunganus

Table 2. Continued.

E. multivirgatus Species Group (Tanner, 1957; Robinson, 1979)
E. mullilineatus (Tanner, 1957; Legler and Webb, 1960)

E. multivirgatus (Lowe, 1955; Mecham, 1957, 1980)

E. brevirostris Species Series

E. brevirostris Species Group (Dixon, 1969; Robinson, 1979)
E. colimensis
E. copei
E. brevirostris
E. dugesii
E. ochoteranae
E. parviauriculatus
E. parvulus

E. egregius Species Group
E. egregius (Mount, 1965, 1968)

E. skiltonianus Species Group (Rodgers and Fitch, 1947)

E. gilberti
E. lagunensis
E. quadrilineatus
E. skiltonianus

* Although Eiselt, and later Mertens, proposed a schneideri “Ras-
senkreis” to include the species listed here as subspecies of a single
form (Eumeces schneiderii), it is now clear from karyological data
(Kupriyanova, 1973; Talliuri, 1975; J.W. Wright, LACM, pers.
comm.) that at least two different karyotypes are involved among
these nominal taxa. The E. schneiderii Species Group is thus in need
of taxonomic re-evaluation with respect to this new information.

into its own section (Section II) and allocated the E. obsoletus
Group to Section III. In my classification, only two sections
are recognized: the Eumeces Section (Taylor’s Section I) and
the Pariocela Section (Taylor’s II and III). Taylor’s placement
of the E. longirostris Group into a separate section arose
from his emphasis upon its unique arrangement of preanal
scales. My combination of the E. longirostris Group with the
E. obsoletus Group is based upon a shared arrangement of
lateral scales that is otherwise unique in the genus. Thirdly,
I retain separate the E. lynxe and E. sumichrasti groups, but
unite them at the species series level. I concur with others
(Smith and Etheridge, 1953; Parker, 1960) that the species
involved are related, but perceive the differences between the
two groups to be of a magnitude greater than that found
within the species group levels in the rest of the genus.

The overall phenetic basis for the erection of the various
categories above the species group level is indicated in the
following key to those groups.

1 a. Median row(s) of dorsal scales conspicuously wider than

adjacent scale rows Eumeces Section ... 2

lb. Median row(s) of dorsal scales not noticeably wider

than scales of adjacent rows

Pariocela Section ... 4

2a. Most of the enlarged median dorsal scales in a single
row at midbody E. taeniolatus Series ... 3

Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus 13

2b. Two rows of enlarged dorsal scales at midbody

E. schneideri Species Group and Series

3a. Two presuboculars posterior to second loreal

E. taeniolatus Species Group

3b. Three presuboculars posterior to second loreal

E. schwartzei Species Group

4a. Inferior lateral body scales in parallel rows 6

4b. Inferior lateral body scales in oblique rows

E. obsoletus Series ... 5

5a. Ground color of juveniles and subadults black, labials
white-spotted; dorsal scales of adults yellow to green-
ish-gray with black scale margins; black scale margins

occasionally expanded into dark body stripes

E. obsoletus Species Group

5b. Juveniles and subadults with distinct dark lateral stripes
and dorsolateral light lines; dorsum of adults bronze or
greenish with black suffusions on the anterior-medial

portions of most dorsal scales

E. longirostris Species Group

6a. A middorsal light line present anteriorly or throughout
body length in juveniles and patterned adults; line ex-
tending anteriorly on head terminating or bifurcating

on the posterior part of the frontal scale

E. lynxe Series ... 7

6b. Middorsa! light line absent in juveniles and patterned
adults; or, if present, terminating or bifurcating on nu-
chal scales 8

7a. Middorsal light line extending posteriorly no more than

a third of the body length

E. lynxe Species Group

7b. Middorsal light line extending posteriorly throughout
body length to tail . . E. sumichrasti Species Group
8a. Scale lying medial to postgenial scale longer than wide,

dorsal median light line present or absent 9

8b. Scale lying medial to postgenial scale wider than long,

dorsal median light line invariably absent 10

9a. Dorsolateral light lines occupying second and third lat-
eral scale rows on the neck just anterior to shoulder

10

9b. Dorsolateral light lines absent, or occupying third and
fourth, or fourth only, lateral scale rows on the neck

region just anterior to shoulder 12

10a. Postnasal scales absent 11

10b. Postnasal scales present

E. skiltonianus Species Group

I la. Supraocular scales four, or if less than four, postmental

scale entire E. brevirostris Species Group

I I b. Supraoculars three, postmental divided by a transverse

suture E. egregius Species Group

12a. Scales around body in more than 24 rows 14

12b. Scales around body 24 or less 13

13a. Keeled lateral postanal scales present

E. fasciatus Species Group, in part

1 3b. Keeled lateral postanal scales absent

E. multivirgatus Species Group, in part

14a. Single dark lateral stripe present on each side of the
body, terminating at the shoulder, at midbody, or on

the tail within three to five scales posterior of vent

E. anthracinus Species Group, in part

14b. Single dark lateral stripe absent or extending through-
out body length and onto tail for a distance consider-
ably more than five scale-lengths posterior to vent . .

15

15a. Postnasal scales present 16

15b. Postnasal scales absent 17

16a. Scales around body 26 13

16b. Scales around body 28 or more

E. fasciatus Species Group, in part

17a. Middorsal light line present in patterned adults and
juveniles, line bifurcating on the nuchals and extending

throughout body length 13

1 7b. Middorsal light line absent, or, if partially expressed,
does not form a bifurcating mark on the nuchals ....

18

18a. Postmental scale divided by a transverse suture and
the light lateral line passes through the ear opening .

E. multivirgatus Species Group, in part

18b. Postmental scale entire, or, if divided, the light lateral

line above the ear opening

E. anthracinus Species Group, in part

SPECIMENS EXAMINED

Eumeces tetragrammus tetragrammus (Specimens exam-
ined, 162). MEXICO. COAHUILA: 3 mi. W Cuatro Cien-
egas (AMNH 77316); 3 mi. NW Cuatro Cienegas (TCWC
40750-51); 12.9 mi. E Cuatro Cienegas (TCWC 40752).
NUEVO LEON: Arroyo de las Vacas, 2 km W Hwy 85 at
Ranchitos (MVZ 185745). QLIERETARO: 9.3 mi. E Jalpan,
3500 ft. (TCWC 29546); El Trapiche (TCWC 45494-97); 1 1
nn. N Jalpan, 2300 ft. (TCWC 32289-90). SAN LUIS PO-
TOSI: 4 mi. (by rd to Oviedo) SSW Ajinche (LSUMZ 2374,
2376); 10 mi. S Antiguo Morelos (FMNH 105277); 1 mi. W
Chantol (TCWC 59971); 3.5 mi. W Chantol, Rancho Pago
Pago (TCWC 59969); Ebano (LSUMZ 343); 7 mi. W El
Naranjo (BCB 6 1 -878); 7 mi. N Valles, Los Sabinos (AMNH
66999); 8 mi. N Valles (UMMZ 118200); 5.4 mi. S, 1.1 mi.
E Valles off Hwy 85 (TCWC 59904). TAMAULIPAS: 0.4
mi. SW Altamira (TNHM 28903-04); 0.3 mi. SW Rancho
Carricitos, 1950 ft. (TCWC 49978); 1.3 mi. WSW Rancho
Carricitos, 2500 ft. (TCWC 49779); 1 mi. E Chamal (UMMZ
101433); 6 mi. NW Chamal (BCB 68-49); 8 mi. NW Chamal
(BCB nh8-908, -909); La Clementina (FMNH 105225); 3.5
mi. WSW Gavilan (TCWC 49780); Gomez Farias (UMMZ
1 10801); 24 mi. SW Jiminez (BCB 3238); Jaumave (UMMZ
95227); 3 mi. NW Limon (BCB 68-44); 19 mi. N Limon,
500 ft. (BCB 7323); 26 km N El Limon (UIMNH 22443);
12 mi. NW Llera (BCB 66-107); 22 mi. SE Manuel (BCB
675 1); Matamoros (USNM 165662, lectotype); Padilla
(TCWC 6937-38); San Jose (UMMZ 69252); Sierra San Car-
los, 1.5 mi. NW Tinaja, 1800 ft. (TCWC 38666-67); Sierra
de Tamaulipas, Hacienda Acuna (UMMZ 101431-32); 18
mi. N Ciudad Victoria (SM 6973); 19 mi. N Cd. Victoria,

14 Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus

Rio Corona (BCB 1 1710); 21 mi. N Cd. Victoria, Rio Corona
(BCB 1 1 7 1 1 ); 22 mi. N Cd. Victoria, Rio Corona (BCB nhv67-
470-nhv67-472); Zaragoza (BCB 68-45). USA. TEXAS:
Cameron Co.: 2 mi. S Bluetown, 50 ft. (BCB 3447-51); 2
mi. W Bluetown, 40 ft. (BCB 4749-5 1 ); Brownsville ( AMNH
102620; FMNH 5499; KU 69025; TAIC 2413 [2]; TCWC
8969-72); Brownsville, Fort Brown (AMNH 79091; USNM
52301); 6 mi. N Brownsville on Fos Fresnos Rd (TNHM
13617-23); 5 mi. SE Brownsville (TAIC 2516); 9 mi. SE
Brownsville (AMNH 79089-90; ANSU 6039); 10 mi. SE
Brownsville, 25 ft. (BCB 3283-86); 17 mi. E Brownsville
(BCB 25); 20 mi. N Brownsville (KU 7754-58); 1 mi. E Los
Fresnos (TNHM 14998-99); 4 mi. W Los Fresnos (TNHM
11924-26); Harlingen (FMNH 94823-24); 1 mi. E Harlin-
gen, 40 ft. (BCB 19); Padre Island (AMNH 8160); 3 mi. SE
Santa Maria, near La Feria Pump Sta. (TNHM 13624);
Southmost Palm Grove (TCWC 38759-60); Duval Co.: Freer
(LSUS 3222); Frio Co.: 1 1 mi. W Dilley (CM 10558); Hi-
dalgo Co.: 5 mi. S Alamo (LSUMZ 18214, 18266); 13 mi.
N Edinburg, La Coma Ranch (TCWC 36524-31); 0.5 mi. N
Hidalgo, FmRd 1962 (TCWC 18176-80); 1 mi. S Hidalgo,
near Rio Grande River (TCWC 1 8 1 81-82); Kenedy Co.: King
Ranch, Norias Div., Rudolf Gate Area (TCWC 38855); Live
Oak Co.: 8 mi. W Jet. FmRd 624 & US Hwy 281 (TAIC
1 17); Starr Co.: Arroyo Los Alamos, 3 mi. SE Rio Grande
City (FMNH 105226); Arroyo El Salado (KU 7747); 6 mi.
W El Sauz (BCB 68-841, 842); Uvalde Co.: 8 mi. N Uvalde
(TCWC 44175, 77176-83); Webb Co.: 0-2 mi. W Bruni,
along RR tracks (TCWC 39270-76, 41 555-6 1; UTEP 8759);
Willacy Co.: Raymondville (TCWC 35558); 3 mi. N Ray-
mondville (MVZ 68402). Other significant localities repre-
sented by unexamined specimens include: TAMAULIPAS:
1 mi. NW La Pesca (Baker and Webb, 1967). VERACRUZ:
Tampico (Taylor, 1935b); 39 mi. and 34 mi. S Tampico
(Darling and Smith, 1954). Specimens examined for which
the locality data are questionable include: MCZ 93177 (Ma-
tegalpa, Nicaragua [sic]); USNM 7858 1 (TEXAS: Rio Grande,
Brule)— I have been unable to locate this placename within
a modem county; WW unnumbered, four specimens (TEX-
AS: San Patricio Co.: Welder Wildlife Refuge, HQ areal-
circumstances surrounding the capture of the specimens (E.
Blacklock, pers. comm.) strongly suggests they were acciden-
tally imported in ornamental vegetation from the lower Rio
Grande Valley.

Eumeces tetragrammus brevilineatus (Specimens exam-
ined, 330). MEXICO. CHIHUAHUA: 5 mi. N Cerro La
Campana (MVZ 70702-03); Santa Clara Canyon, 4.5 mi. (by
rd) E MX Hwy 45 (LACM 1 16401); Sierra del Nido, 4.7 mi.
(by rd) W Encinillas (UTEP 62). COAHUILA: 21 mi. NW
Ciudad Melchor Muzquiz (EAL 3139); 16 mi. E, 18 mi. N
Ocampo (KU 38073); 5 mi. W Piedra Blanca, 5000 ft. (MVZ
58338); 5.3 mi. E Piedra Blanca (UAZ 16815-17); 8 mi. SW
Piedra Blanca, 7000 ft. (MVZ 58337); Sierra Madera Jardin
(SRSU 864); 2 mi. S Villa Acuna (UIMNH 27 1 36); NUEVO
LEON: near Sabinas Hidalgo (UIMNH 2244 1 ). USA. TEX-
AS: Atascosa Co.: 4 mi. W Jordanton (USL 15433); near
Lytle (KU 15564); Bandera Co.: 9 mi. S Medina (TCWC

1 5065); 7 mi. SW Medina on W Fork Medina River (TNHM
1410); 18 mi. NW Medina, Sutton’s Ranch (TNHM 1898);
8.4 mi. N Vanderpool on Hwy 187 (LSUMZ 10359); 10.4
mi. W Vanderpool, Hwy 337 (USL 13133); 14.7 mi. W Jet.
Hwy 462 on Hwy 470 (USL 13133, 13551, 15680); Bell Co.:

7.2 mi. NE Holland along Salado Creek (TCWC 23064);
Bexar Co.: Helotes (KU 7744, 7764; USNM 10527 [2],
1 3628); Marnock’s Ranch, near Helotes (KU 690 1 8; USNM
10159-B, lectotype, and 10159-A, syntype); 1 mi. N Helotes
(MVZ 68397); 7 mi. SE Lytle (CM 18406); 6 mi. NW Rio
Medina (CM 58465); San Antonio (KU 8703, 8810-11,
I 5565); San Antonio, Brackenridge Park (CM 1 8392); 9 mi.

5 San Antonio (CM 8466); 8 km ENE Shavano Park, Voight
Ranch (UTEP 8873); Somerset (KU 8703); 2.5 mi. N Som-
erset (BCB 2257); 6 mi. N Somerset, Medina River (CM
18425); 8 mi. SW Somerset (BCB 3351); Von Ormy (CM
1 8377); Blanco Co.: 4.5 mi. SE Johnson City (TCWC 8942);

6 mi. S and 3.4 mi. E Johnson City (UTEP 165); Bosque
Co.: 2 mi. S Mosheim, Mid Fork Bosque River (TCWC
36932, 38740-43); 3 mi. N Osage (TCWC 14242); Brewster
Co.: Alpine (SRSU 223, 343); 3 mi. SW Alpine (KU 7768-
70); 20 mi. S Alpine (SRSU 453); Big Bend Natl. Park (BBNP),
Boquillas Ranger Station (UNM 9974); BBNP, base of Burro
Mesa (UNM 9972-73); BBNP, Casa Grande (TCWC 1 6052);
BBNP, Government Springs (UNM 20843); BBNP, Gov-
ernment Wells (UNM 1 8239-40); BBNP, Grapevine Springs
(TCWC 16045); BBNP, south of Moss Well, 5000 ft. (UNM
5900); BBNP, Oak Canyon, 4000 ft. (UNM 20846); BBNP,
Panther Jet. (UNM 6555); Chisos Mts., E slope (KU 1 3200);
Chisos Mts., Basin, 6000 ft. (TCWC 1113); Chisos Mts.,
Green Gulch (TCWC 16050); Chisos Mts., Mt. Emory, 8000
ft. (KU 12748); Chisos Mts., Pine Canyon (TCWC 14269);

16.2 mi. NW La Linda, Coahuila (EAL 3238); 60 mi. S
Marathon, Black Gap Wildlife Mgmt. Area (TCWC 20161;
TNHM 12939, 12987); Glass Mts., 5 mi. N Marathon (KU
13199); Strumbere Ranch (SRSU 3605); Brown Co.: 4 mi.
W Bangs, US Hwy 67 (TCWC 23446-48); 4 mi. W Brown-
wood (KU 1 1 387); Burnet Co.: “Burnet County” [no further
data] (USNM 5877); Burnet (CAS 7409); 8 mi. W Burnet
(TCWC 4499-501); Callahan Co.: 19 mi. W Cross Plains
(TNHM 9738); Comal Co.: New Braunfels, Water Rec. Dist.
#2 (TCWC 1 5064); Comanche Co.: 5 mi. N DeLeon (TCWC
1 5266); Crockett Co.: 1 1 mi. E Ozona, US Hwy 290 (ANSU
148); Dimmit Co.: near Carrizo Springs, Nueces River (KU
8195-96); Edwards Co.: 24 mi. NE Rocksprings (TCWC
4502); Gillespie Co.: 15 mi. NE Fredricksburg(TCWC 5545-
46); Hays Co.: Fern Bank Springs, Little Arkansas (TCWC
31486-87, 36534, 38744); 10 mi. S Oak Hill on Big Bear
Creek (TNHM 21179); Pollard Wildlife Refuge (TCWC
38745-49); 4 mi. W San Marcos (TCWC 8941); 7 mi. W
San Marcos (FSM 3579); 2 mi. E Wimberly on Cypress Creek
(TNHM 8794); 4 mi. E Wimberly (TCWC 27320-24); Irion
Co.: 1 1 mi. NW Mertzon (ANSU 2768-69, 277 1 , 2773, 2790);
12 mi. NW Mertzon (ANSU 4535); 13 mi. NW Mertzon
(ANSU 629-30, 871-73); Jeff Davis Co.: Cherry Valley (MCZ
12822); 10.6 mi. N Fort Davis, Jones Ranch (TCWC 26108);
Nations Canyon (SRSU 688); Kendall Co.: 0.5 mi. W Cen-

Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus 15

tury Caverns (TCWC 38739); 0. 5-1.0 mi. S Camp Alzafar
(TCWC 30172, 38737-38); Kerr Co.: 4 mi. NE Centerpoint
(TCWC 181); Kerrville (USL 1716); 15 mi. SE Kerrville
(LSU 13676); Kerr Wildlife Mgmt. Area (TCWC 13806);
Kimble Co.: 2-4 mi. N Cleo (TCWC 38750-53); 2 mi. SW
Junction on US Elwy 377 (UTEP 8583); 10 mi. W Junction
(ANSU 7153); 15 mi. NW Junction (ANSU 8132); 2.3 mi.

5 Junction, Schriener Ranch (TNHM 7065); Telegraph, Point
Creek and Schiener Ranches (TNHM 7003-05); Kinney Co.:
Ft. Clark (USNM 25445); Live Oak Co.: 6 mi. E George
West on Nueces River (TAIC 258.1-.2); 3 mi. W Three
Rivers (TCWC 10535-36, 10538); 8 mi. W Jet. FmRd 624

6 US Hwy 281 (TAIC 123.2-.3); Mason Co.: 20 mi. ESE
Mason (TCWC 31047-50); 8 mi. S Mason (TCWC 3105);
10 mi. S Mason (TCWC 31052-53; Maverick Co.: Mangus
Ranch, S of Eagle Pass (TAIC 2473, [2]); McClennan Co.:
Bluff Creek, 0.5-2 mi. W Crawford (FMNH 46757-58,
46762); near Crawford (FMNH 46759); McMullen Co.: 5.6
mi. W Whitsett (TCWC 39267); 7.9 mi. W Whitsett (TCWC
39266); Medina Co.: “Medina County” [no further data]
(USNM 42307); 5 mi. N Hondo (TCWC 14621); 3 km ENE
Mico, 1200 ft. (UTEP 9481); Nueces Co.: 10 mi. W Corpus
Christi (TCWC 18175); Palo Pinto Co.: BSA Camp Con-
stantine (TCWC 25271); 10 mi. W Graford (ANSU 7154);
10 mi. S Mineral Wells (TCWC 406); 2.8 mi. N Palo Pinto
(TNHM 1 3503); Pecos Co.: 1 1 mi. E Bakersfield (SRSU 77 1 );
Fort Stockton (SRSU 1412); near Iraan (TNHM 33380); 1.4
mi. W Sheffield on US Hwy 290 (UTEP 8584); Presidio Co.:
Chinati Mts., Pinto Canyon (SRSU 524); 1 1 mi. W Valentine,
CE Miller Ranch (TNHM 1 124, 2876, 3257, 3298-99, 3395,
4256, 4283); Regan Co.: Best (ANSU 2745-47); 15 mi. W
Big Lake (ANSU 528); 9 mi. W, 12 mi. S Big Lake, 2500 ft.
(ANSU 7166-69, TCWC 31355-58); Real Co.: 14.4 mi. E
Jet. US Hwy 83 on FmRd 337 (USL 16378); San Saba Co.:
Gorman Falls Camp, 6 mi. SE Bend (ANSU 141, 144, 2799-
800, 5525, 5629, 5264); San Saba (ANSU 2916, 4064-74);
1 mi. S San Saba (ANSU 2968); 12 mi. E San Saba (TNHM
28837); 20 mi. NNW San Saba (TNHM 9697); Schleicher
Co.: “Schleicher County” [no further data] (ANSU 2729); 5
mi. S Christoval (ANSU 145); 12 mi. S Christoval (ANSU
146); Shackleford Co.: 20 mi. N Albany, Matthews Ranch
(UNM 9049); Sutton Co.: 5 mi. S Sonora (TNHM 33381);
1 0 mi. S Sonora ( M VZ 38 1 99-200); 26 mi. SE Sonora (USNM
6529); Terrell Co.: 27 mi. ESE Dryden (USL 15432); 13 mi.
S Sheffield (TNHM 7588, 7612, 7678, 7777); 15 mi. S Shef-
field, Blackstone Ranch (TNHM 7105); 18 mi. S Sheffield
(TNHM 8131-33); 21 mi. S Sheffield (TNHM 7920, 7948);
30 mi. S Sheffield, Chandler Ranch (SRSU 303, 316, 325,
338-40, 2201; TNHM 8314-18, 8054); Tom Greene Co.:
Christoval (ANSU 1 1 80, 2108-10, 5579); 3 mi. E Christoval,
Toe Nail Trail (ANSU 140, 1 42); 4 mi. S Christoval, Concho
River (ANSU 5561); 4.5 mi. S Christoval, Head of River
Ranch (ANSU 147, 150); 5 mi. S Christoval (ANSU 149);
N Concho Lake (ANSU 279, 3114-15, 3161); Fort Concho
(USNM 1 2777); 4 mi. N Log Cabin Steakhouse Club (ANSU
5274); Nasworthy Lake (ANSU 3035-36); San Angelo (ANSU
3 1 40, 8 1 3 1 ); 4 mi. S San Angelo, near Nasworthy Dam (ANSU
143); Travis Co.: Austin, Texas Univ. Campus (TNHM

25727); 1 mi. NW Austin, Bull Creek Rd (TNHM 33375);
4 mi. SSW Austin (TNHM 9877); 5 mi. SW Austin, Barton
Creek (TNHM 1693, 1936, 5870, 5942); 6 mi. SE Austin
(TNHM 32 1 95); 1 .5 mi. SE Manchaca (TNHM 2 1 683; Shoal
Creek (TNHM 13074-75); 1 mi. upriver from Zilker Park
(TNHM 13501); Uvalde Co.: ne&r Concan, Bludworth Ranch
(SM 5024); 3 mi. E Concan (BCB 7161); 3 mi. N Sabinal
(TNHM 4873); 8 mi. N Uvalde (TCWC 44173-74); Val
Verde Co.: 3 mi. W Comstock, along RR (ANSU 7161); 5.2
mi. N Comstock at Pecos River (ANSU 7156); 12 mi. N
Comstock at Pecos River (ANSU 7 1 57-60); 1 2 mi. S, 1 1 mi.
E Comstock (ANSU 7165); 19.5 mi. SE Comstock (ANSU
7 1 55); 20 mi. SE Comstock, under US Hwy 90 (ANSU 7 1 62—
64); near mouth of Devils River (KU 7748); Dolan Falls,
Devils River (UNM 5973); Dolan Springs, Fawcett Ranch
(UNM 5971-72); 12 mi. NW Del Rio (TNHM 32495); 50
mi. NW Del Rio (MVZ 68398); 60 mi. SW Ozona (ANSU
5448); Webb Co.: Laredo (UMMZ 1 14253); Wilson Co.:
Cibolo River bottoms (SM 4353, 4355). Other significant
localities represented by unexamined specimens include:
MEXICO. COAHUILA: 33 mi. N, 8 mi. W San Geronimo
(KU 33502); NUEVO LEON: 4 mi. W Sabinas Hidalgo (Tay-
lor, 1935b); 31 mi. S Sabinas Hidalgo (Taylor, 1935b). TEX-
AS: Coke Co.: 2 mi. S Blackwell (Brown, 1950); Coleman
Co.: 22 mi. S Valera, Day Ranch (TCWC 53456-61); Kinney
Co.: 18.0 mi. N Bracketville (TCWC 46527); Llano Co.: 19
mi. SW Llano, FmRd 2323 (TCWC 58480-81); McCulloch
Co.: 8 mi. N Brady (TCWC 18925); Menard Co.: 2.5 mi. E
Ft. McKavett, 1800 ft. (TCWC 51 196); Throckmorton Co.:
19 mi. NW Albany (KU 61796-97). Specimens examined
for which the locality data are questionable include: FMNH
106623-26 (“large spring near Sabinas, Hidalgo”)— the lo-
cality may refer to Sabinas Hidalgo in Nuevo Leon; TNHM
27810 (OKLAHOMA; 5 mi. SW Colbert); FMNH 27215-
1 7 (“Probably Brownsville, Texas”); BCB nH69- 1237,-2181
(TEXAS: Henderson Co.: 10 mi. S Athens); FMNH 46760-
6 1 (TEXAS: Limestone Co.: between Oletha and Thronton).

Eumeces tetragrammus tetragrammus x brevilineatus
(Specimens examined, 10). MEXICO. NUEVO LEON: Ar-
royo de las Vacas, 2 km W Hwy 85 at Ranchitos (MVZ
185746). USA. TEXAS: Jim Wells Co.: Casablanca, Nueces
River (KU 8812); La Salle Co.: 2.8 mi W Jet. FmRd 624,
FmRd 468, and FmRd 469 (TAIC 643); Live Oak Co.: 3 mi.
W Three Rivers (TCWC 10537); 8 mi. W Jet. FmRd 624
and US Hwy 281 (TAIC 123.1); McMullen Co.: 17 mi. S
Charlotte (TNHM 28836); 6.9 mi. S Tilden on State Hwy
16 (LACM 1 34855); 8 mi. W Whitsett (TCWC 39265); Uvalde
Co.: 8 mi. N Uvalde (TCWC 44171-72).

Eumeces tetragrammus callicephalus (Specimens exam-
ined, 106). MEXICO. CHIHUAHUA: Bavispe River below
Three Rivers, Sonora-Chihuahua line (BYU 13145-49,
14233); ca. 2 mi. E Cerocahui (BYU 14248-49); Cuiteco
(BYU 14259-61, 14608-10); Guasaremos (MCZ 43389-90);
Madera (MCZ 15928); Madronoi, W Rim, between Urique
and Cerocahui (BYU 14338); 8 mi. W Matachic (AMNH
68295); Pacheco (MVZ 46672); 3 mi. NE Temoris (KU
51462). JALISCO: 38.2 mi. NW Guadalajara on MX Hwy
15 (FSM 12844-45); Hostotipaquillo (AMNH 17943); near

16 Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus

Magdalena (FMNH 106357-58); 3 mi. NE Magdalena (KU
38069). NAYARIT: 6 mi. SE Ahucatlan (UMMZ 1 18530);
7.1 mi. N Compostela, 3200 ft. (FSM 28999); Rosamorada
(AMNH 1 5488); Mt. San Juan Tepee (FMNF1 1 06359); La-
guna Santa Maria del Oro, 2350 ft. (AMNH 96608); 19.6
mi. E Santa Cruz (CAS 95518); Santiago Ixcuintla (AMNH
19305-06); 12 mi. SW Santiago Ixcuintla (FSM 19305-06);
5 mi. NE Sentispac (AMNH 87673-77, 87974); 23. 1 mi. E
Tepic (MVZ 71259); 29 mi. SW Tepic at Crater Lake (BCB
64-1331). SINALOA: 16 km NNE Choix, 1700 ft. (KU
73745); vie. El Dorado (BCB 66-1381); La Cruz (LACM
6768); 1 mi. N Mazatlan (UTEP 4863-65); 2 mi. E Mazatlan
(CAS 104975-77); 5 mi. N Mazatlan, along Sabalo Beach
Rd (AMNH 87672); Sierra Surutato, 0.5 mi. (by rd) SE Los
Hornos, ca. 1920 m (CAS 155910, 155913); Sierra Surutato,
Canon de Tarahumare between La Joya and Baranca de las
Tahonitas, ca. 1310m (CAS 15591 1-12); Teacapan (LACM
6769-70); 24.8 mi. (by MX Hwy 40) E Jet. MX Hwy 15
(CAS 11483). SONORA: Alamos (AMNH 64219; UAZ
3468); 7 mi. SE Alamos, Arroyo Cuchujaqui (MVZ 72602);
N Slope Alamos Mt., Las Higueras Creek (UAZ 3472); stream
above Alamos on Sonora-Chihuahua border (MVZ 74186);
Arispe, 840 m (UAZ 3471, 3473-74); Guirocoba, Cienigitas
(MVZ 50735); 9 mi. NNE Imuris, 1000 m (KU 50633); 9.4
mi. by rd from Huertas (UAZ 1 1310-1 1); ca. 28 mi. E Na-
cozari, Presa de Rebeico (UAZ 1 1309); 36 mi. ENE Nuri,
3600 ft. (UAZ 3467); upper fork Nutria Creek (BYU 1 3 1 40-
41); 2 mi. E Santa Ana on rd to Yecora (UAZ 28193-94);
above Santa Maria Mine (UMMZ 78124); 0. 5-1.0 mi. SW
Yecora (UAZ 16598); Rio Zatachi, 2930 ft. (UAZ 3475).
USA. ARIZONA: Cochise Co.: Huachuca Mts. (CAS 48095-
96, 80747); Huachuca Mts., Ash Canyon (KU 6473-76);
Huachuca Mts., Copper Canyon (FMNH 46 1 1 7); Huachuca
Mts., Hunter Canyon, 5640 ft. (UAZ 1 6734); Huachuca Mts.,
Ramsay Canyon (UMMZ 71029); N Ridge Hunter Canyon,
Short Springs (UAZ 16737); Pima Co.: Baboquivari Mts.,
Brown Canyon (AMNH 86573); Baboquivari Mts., Elkhorn
Ranch, 3750 ft. (UAZ 7166); Santa Cruz Co.: Pajarito Mts.,
Pena Blanca Canyon (AMNH 15063); Pajarito Mts., Pena
Blanca Springs (MVZ 49838, 53877; CAS 84125); Pajarito
Mts., Walker Canyon, 1.5 mi. N Ruby Rd, Rockwell Camp,
3750 ft. (UAZ 19834, 30607-08); Santa Rita Mts., Madera
Canyon (AMNH 64342; LACM 5929); Sycamore Canyon,
3800-3960 ft. (UAZ 3466, 3419; MVZ 50736). NEW MEX-
ICO: Hidalgo Co.: Guadalupe Mts., Guadalupe Canyon, 4500
ft. (KU 74332-34). A significant locality represented by an
unexamined specimen is: MEXICO. ZACATECAS: Mes-
quital del Oro (Taylor, 1935b). Specimens examined for which
the locality data are questionable include: ANSP 13604-05
(MEXICO. GUANAJUATO: Guanajuato).

ACKNOWLEDGMENTS

I thank the following for allowing me to examine specimens
in their care: R.G. Zweifel, American Museum of Natural
History (AMNH); E.V. Malnate, Academy of Natural Sci-
ences of Philadelphia (ANSP); G.G. Raun, San Angelo State
University (ANSU); .1. Sites and W.W. Tanner, Brigham

Young University (BYU); A.E. Leviton, California Academy
of Sciences (CAS); C.J. McCoy, Carnegie Museum (CM);
E.A. Liner, E.A. Liner Collection (EAL); H. Marx, Field
Museum of Natural History (FMNH); H.W. Campbell, Flor-
ida State Museum (FSM); W.E. Duellman, University of
Kansas Museum of Natural History (KU); J.W. Wright, Nat-
ural History Museum of Los Angeles County (LACM); D.A.
Rossman, Louisiana State University Museum of Zoology
(LSUMZ); L.M. Hardy, Louisiana State University, Shreve-
port (LSUS); E.E. Williams, Museum of Comparative Zo-
ology (MCZ); H.W. Greene and R.C. Stebbins, Museum of
Vertebrate Zoology (MVZ); B.C. Brown, Strecker Museum
(SM) and B.C. Brown Collection (BCB); J.F. Scudday, Sul
Ross State University (SRSU); A.H. Chaney, Texas A&I
University (TAIC); J.R. Dixon, Texas Cooperative Wildlife
Collection (TCWC); R.F. Martin, University of Texas Nat-
ural History Museum (TNHM); C.H. Lowe, University of
Arizona (UAZ); L.E. Maxson, University of Illinois Natural
History Museum (UIMNH); C.F. Walker, University of
Michigan Museum of Zoology (UMMZ); W.G. Degenhardt,
University of New Mexico Museum ofSouthwestern Biology
(UNM); E.D. Keiser, University ofSouthwestern Louisiana
(USL); J.A. Peters, National Museum of Natural History
(USNM); R.G. Webb, Laboratory for Environmental Biol-
ogy, University of Texas at El Paso (UTEP); E. Blacklock,
Welder Wildlife Refuge (WW). Assistance in the field was
provided by Douglas Albaugh, James C. Kroll, Tearil W.
Lewis, Val .1. Roessling, Jr., and Robert A. Thomas. In ad-
dition, Robert L. Bezy, James E. DeWeese, D.F. Hoffmeister,
John P. Kargcs, David Lintz, Richard B. Loomis, David J.
Morafka, Robert W. Murphy, Hobart M. Smith, and Robert
A. Thomas were especially helpful in providing museum
data, literature, or unpublished information on these skmks.
James R. Dixon and John W. Wright encouraged me to
publish this study, even though the latter has continuing
doubts about its conclusions. Robert Bezy, Robert G. Webb,
and John W. Wright read and edited early versions of the
manuscript.

Photographic assistance and materials were provided by
several individuals, notably Richard J. Baldauf, Robert L.
Bezy, Tearil W. Lewis, and Robert A. Thomas. Collections
of lizards in Mexico during 1971 were made possible by
permits issued by the Direccion de la Fauna Silvestre of the
Republic of Mexico.

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Accepted 17 July 1984.

Contributions in Science, Number 357

Lieb: Skinks Allied to Eumeces tetragrammus 19

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Number 358
11 April 1985

CONTRIBUTIONS IN

SYSTEMATICS OF THE SOUTH AMERICAN FRESHWATER
FISH GENUS ADONTOSTERNARCHUS
(GYMNOTIFORMES, APTERONO TIDAE)

Francisco Mago-Leccia, John G. Lund berg, and

Jonathan N. Baskin

- J.

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SYSTEMATICS OF THE SOUTH AMERICAN FRESHWATER
FISH GENUS ADONTOSTERNARCHUS
(GYMNOTIFORMES, APTERONOTIDAE)

Francisco Mago-Leccia,1 John G. Lundberg,2 and
Jonathan N. Baskin3

ABSTRACT. The apteronotid fish genus Adontosternarchus Ellis,
1912, is revised, and characters are presented as evidence for the
monophyly of the genus and of the interrelationships of its member
species. Four species are recognized of which two are described as
new: A. sachsi (Peters, 1877), A. balaenops (Cope, 1878), A. deve-
nanzii new species, and A. clarkae new species. The known distri-
bution of each species is plotted and a key to species is provided.
Recent collections in channels of the lower Orinoco River show A.
sachsi and A. devenanzii to be extremely abundant.

RESUMEN. Este trabajo es una revision sistematica del genero
apteronotido Adontosternarchus Ellis, 1912, e incluye la presenta-
tion de caracteres que evidencian su condition monofiletica, asi
como tambien una hipotesis explicita de interrelaciones entre sus
especies integrantes. Se reconocen cuatro especies en el genero, de
las cuales dos son nuevas para la ciencia: A. sachsi (Peters, 1877),
A. balaenops (Cope, 1878), A. devenanzii sp. n. y A. clarkae sp. n.
La distribution geografica conocida para cada especie se muestra en
mapas. Asi mismo, se senala que colecciones recientes hechas en los
canales del Bajo Orinoco revelaron la extraordinaria abundancia de
las especies A. sachsi y A. devenanzii.

INTRODUCTION

The genus Adontosternarchus was established by Ellis (1912:
424) to distinguish apteronotids characterized by “Teeth
wanting; lower jaw with a distinct V-shaped median groove
for the reception of the pointed decurved upper jaw.” The
type species, by monotypy, and subsequent designation, is
Sternarchus sachsi. a species described by Peters (1877) based
on specimens collected by Dr. Carl Sachs from the Vene-
zuelan llanos, near San Fernando de Apure. Ellis referred to
A. sachsi all material of Adontosternarchus available to him
from the Amazon basin. In 1 942 Eigenmann and Allen added
Adontosternarchus balaenops (Cope), a species based on a
single poorly preserved specimen from Peru.

Recently we made large collections of Adontosternarchus
from the Orinoco Delta and middle Orinoco that disclosed
the presence there of two species. Further study demonstrated

Contributions in Science, Number 358, pp. 1-19
Natural History Museum of Los Angeles County, 1985

a basis for recognizing four species in the genus. Only part
of the material treated by Ellis represents A. sachsi. The fish
he illustrated (1913: pi. 22, fig. 3; see our Fig. 9) as A. sachsi,
and others in the same series, are A. balaenops, although he
referred the name balaenops to Sternarchella. We redescribe
A. sachsi and A. balaenops, and describe two additional species
as new. One of these is the common middle Orinocan form
modern workers have called A. sachsi (Mago-Leccia, 1967,
fig. 10; 1970). The other was discovered among specimens
provided to us by Ms. Kate Clark from the Rio Negro, Ven-
ezuela, near the Brazilian border, and now is known to be
widespread in the Amazon system.

METHODS AND MATERIALS

Specimens, carefully straightened and pinned down, were
measured with Helios dial calipers. Head measurements were
made under low power magnification. Gymnotiform fishes
often suffer damage (predation) to the tail region. Most
wounded fish are recognized by their truncated, or abruptly
narrowed and/or abruptly depigmented tails, but some in-
dividuals often remarkably regenerate the lost tail and fins.
Despite careful external examination, apparently nearly com-
plete regeneration occurred in some individuals in our mea-
sured samples. These fish are recognized as outliers on the
low ends of the scales for measurements involving the tail
but otherwise they have near average measurements. This
problematic element of morphometric variation makes dif-
ficult both the preparation and the use of identification keys.
Persons working with gymnotiform fishes should be aware
of cryptically damaged but partly regenerated fish. The di-

1. Instituto de Zoologia Tropical, Facultad de Ciencias, Univer-
sidad Central de Venezuela, Caracas, Venezuela.

2. Department of Zoology, Duke University, Durham, North
Carolina 27706, USA.

3. Department of Biological Sciences, California State Polytech-
nic University, Pomona, California 91768, USA.

ISSN 0459-8113

agnostic measurements are: Total length (abbreviated TL),
length from snout to end of base of anal hn (abbreviated
LEA), length of anal fin base, distance from tip of snout to
origin of dorsal thong (a fleshy, ray-less filament attached to
back over about last third of anal fin and the tail base; minor
dissection is often necessary to locate thong’s origin), length
of tail (starting from posterior end of anal fin base), length
of caudal peduncle, distance from snout to origin of anal fin,
snout to vent distance, greatest body depth, head length (tak-
en to upper end of soft opercular membrane), eye diameter,
distance from snout tip to rictus, and size of branchial open-
ing.

Counts of anal, caudal, and pectoral fin rays were taken
with strong transmitted light and include all elements. Ver-
tebral counts (all from radiographs) begin with the first free
vertebra behind the compound Weberian complex. This is
the first one bearing a full neural spine. The last one counted
has its hemal spine immediately behind the base of the last
anal fin ray.

Abbreviations used for various institutions are: Academy
of Natural Sciences, Philadelphia (ANSP), American Mu-
seum of Natural History (AMNH), California Academy of
Sciences (CAS), Carnegie Museum (CM, material now at
FMNH), Duke University (DU), Field Museum of Natural
History (FMNH), Ministerio de Agricultura y Cria, Estacion
de Puerto Ayacucho, Venezuela (MAC-PAY), Museo de
Biologia de la Universidad Central de Venezuela (MBUCV),
Museum of Comparative Zoology, Harvard University
(MCZ), Museu de Zoologia da Universidade de Sao Paulo,
Brazil (MZUSP), National Museum of Natural History,
Smithsonian Institution (USNM), Natural History Museum
of Los Angeles County (LACM), University of Michigan
Museum of Zoology (UMMZ), and Berlin Museum (ZMB).

Adontosternarchus Ellis

Adontosternarchus Ellis, 1912:424 (in Ellis, 1912, type

species: Sternarchus sachsi Peters, 1877, designated sub-
sequently in Ellis, 1913:155 by monotypy).

The valid name Adontosternarchus was first published by
Ellis ( 1 9 1 2) in Eigenmann’s monograph on the freshwater fish-
es of British Guiana. The name and diagnostic characters
appeared just once in a key to genera said to inhabit Guianan
waters. No Adontosternarchus species has been recorded from
British Guiana. In his 1913 revision of gymnotiform fishes
Ellis provided what he clearly intended to be the original
description of this genus and discussion of the only recog-
nized species, A. sachsi.

DIAGNOSIS. Apteronotid fishes with a unique beak-like,
terminal mouth in which the margin of lower jaw is strongly
curved to form a V-shaped notch across the mandibular
symphysis flanked by elevated flanges and, in turn, the snout
is curved downward into notch of mandibles, and the margin
of the upper jaw is concave to received the lower. Other
diagnostic features are the absence of teeth at least in indi-
viduals over about 30 mm TL, and the slightly to markedly
bulbous chin.

DESCRIPTION. Body compressed and moderately elon-

gate; dorsal profile of body nearly straight to gently convex;
ventral profile of body strongly convex to nearly angular at
anal fin origin, nearly straight behind; anal fin origin about
under branchial opening; lateral line complete.

Head small, slightly compressed to rounded; its dorsal pro-
file variable, ventral profile nearly straight; mouth small;
rictus in advance of center or margin of small eye; chin round-
ed to bulbous and projecting. The margin of lower jaw is
strongly curved to form a V-shaped notch across the man-
dibular symphysis flanked by elevated flanges and, in turn,
the snout is curved downward into notch of mandibles and
the margin of the upper jaw is concave to receive the lower.

Eye small; anterior nostril located about midway between
tip of snout and anterior margin of eye, opening at end of a
short tube; posterior nostril without a tube and located above
and slightly in front of anterodorsal margin of eye; branchial
membranes joined to isthmus; branchial opening restricted
to a short oblique slit in front of base of pectoral fin; anus
and short urogenital papilla (both sexes) adjacent, located in
large adult fish between raised rims of united branchial mem-
branes, their positions shift relatively forward with growth
(Fig. 12).

Anal fin elongate with 135-185 rays (Table 2); caudal and
pectoral fins small with 10-22 rays and 12-18 rays respec-
tively (Table 1); body and base of caudal fin scaled; head, fin
membranes, and dorsal thong lacking scales; 4-8 rows of
enlarged cycloid scales along the flanks, including the pored
lateral line scale row; small scales above large scales to dorsal
midline, below to anal fin base and onto breast; scales of
lower flanks with free ventral or ventroposterior margins,
other scales with free posterior margins.

Salient osteological features of Adontosternarchus are: pre-
maxillary bone small and connected to maxillary by a long
ligament; maxillary bone elongate (Figs. 2 A, 3); infraorbital
series represented only by bony, superficial tubes; supratem-
poral and pterotic canals of the laterosensory system of the
head represented by free bony tubes (Fig. 3); posttemporal
fossae absent; cranial fontanelles present, the interfrontal
shorter and broader than the interparietal; lateral ethmoids
and vomer present; mesopterygoid bone short, edentulous
and with a well-developed, ascending process which articu-
lates with orbitosphenoid; preopercular bone broad (Fig. 3);
pectoral girdle without mesocoracoid (Fig. 6); cleithrunt broad;
scapular foramen absent; coracoid with a long ventral process
which fails to reach the cleithral symphysis; posttemporal
fused to supracleithrum; 4 pectoral radials; 5 branchiostegal
rays, the last two greatly broadened, the three anterior more
slender (Fig. 4); urohyal small; gill-rakers reduced to small
bony nodules covered by cartilage; 3 infrapharyngobranchi-
als, the posteriormost one cartilaginous; 5 epibranchials, the
fifth one cartilaginous; upper pharyngeal tooth plate present,
strongly connected by a ligament to epibranchial 3 (Fig. 5);
53-64 vertebrae to base of last anal fin ray (Table 4); We-
berian apparatus without claustrum.

ETYMOLOGY. Greek a = without + odons = tooth +
sternon = breast + archos = anus. Gender masculine.

MONOPHYLY AND RELATIONSHIPS. It is not our
purpose in this paper to accomplish a phylogenetic analysis

2 Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus

Figure 1. Head profiles of Adontosternarchus species. A, A. balaenops ; B, A. devenanzii; C, A. clarkae ; D, A. sachsi. Dotted line indicates
position of anus.

of the Apteronotidae. One of us (FML) is engaged in a study
of that broader subject. However, it is important for future
phylogenetic work to point out the evidence for the mono-
phyly of the genus Adontosternarchus. The hypothesis of ex-
clusive common ancestry of the four species of Adontoster-
narchus is supported by the two features used by Ellis to erect
the genus, i.e., (1) the form of the beak-like snout, bulbous
chin and curved mouth (Fig. 1), and (2) the much reduced
dentition (Figs. 2, 3). The odd form of the chin is due to the
presence of an accessory electric organ formed from sensory
nerve fibers (Bennett, 1971). Additionally, we mention the

small size of the upper jaw elements, and the elongate pre-
maxillary-maxillary ligament (Figs. 2, 3). Based upon com-
parisons with other apteronotids and gymnotiforms these
characteristics appear to be uniquely shared by the species
of Adontosternarchus. Of these features only the nearly com-
plete absence of teeth (present only in juveniles) is ap-
proached by some other apteronotids (e.g., Sternarchogiton
and an undescribed form from the lower Orinoco have lost
upper jaw teeth but retain dentary teeth) but these taxa do
not present facial orgnathal similarities to Adontosternarchus
which can be interpreted as synapomorphies.

Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus 3

Figure 2. Snout region of Adontosternarchus devenanzii sp. n. 1 20.0
mm TL, MBUCV-V-4772. A, lateral view; B, dorsal view; C, ventral
view. EL lateral ethmoid, ET ethmoid (=mesethmoid), FR frontal,
MX maxillary, NA nasal, PAE anterior piece of ethmoid, PAS para-
sphenoid, PMX premaxillary, VO vomer.

Certain aspects of morphometric and coloration diversity
among the species of Adontosternarchus suggest the following
hypothesis of interrelationships. A. sachsi is taken to be the
sister taxon of the three species A. balaenops, A. devenanzii,
and A. clarkae which share a distinctive, boldly mottled color

pattern on the back and sides (Figs. 7, 8, 14, 16). The pig-
mented blotches of this pattern are irregular in outline and
generally cover areas larger than a single scale. A. sachsi has
a nearly uniform coloration of the sides (Fig. 19) although
the scale margins are often darker than their centers. This
uniform pattern appears certainly to be the primitive con-
dition in Adontosternarchus based on outgroup comparison
to other apteronotids ( Apteronotus , Porotergus, Sternarchel-
la, Sternarchogiton, Sternarchorhamphus, and Sternarcho-
rhynchus). The relatively deep body form and short tails of
A. balaenops and A. devenanzii (Figs. 10, 11 and species
diagnoses) are considered shared derived similarities based
on outgroup comparisons; thus these are hypothesized to be
sister species. A. clarkae and A. sachsi are shallower in re-
lation to length and their tails are relatively long, more like
the other apteronotids examined. A. balaenops and A. de-
venanzii each have their own phyletically advanced color
pattern element. A. balaenops possesses black anal and pec-
toral hn membranes. A. devenanzii has a narrow pale or
yellow stripe along the midline from the chin or snout to the
base of the dorsal thong. Other Adontosternarchus, most oth-
er apteronotids (some species of Apteronotus and Sternar-
chorhamphus have black fin membranes; Apteronotus albi-
frons and Sternarchorhynchus curvirostris have broad, light,
mid-dorsal stripes), sternopygids and rhamphichthyids lack
these species-specific novelties. At this time we have not
identified uniquely derived character states for either A. clar-
kae or A. sachsi.

In the context of our hypothesis on species interrelation-
ships and their geographic ranges, A. balaenops (central Am-
azon)and,4. devenanzii (middle and lower Orinoco) are sister
species which arose in allopatry. Because Adontosternarchus
is not present in the Guianas it seems likely that the Casi-
quaire served as the dispersal route for the common ancestor
of balaenops and devenanzii between the Orinoco and Am-
azon although the basin of its origin is uncertain. Collection
records suggest that these species are restricted now to low-
land large rivers and lagoons, and perhaps the steeper gra-
dients of the upper Orinoco mitigate against secondary con-
tact.

A. clarkae is widespread in the upper parts of major Am-
azon tributaries. This species is sympatric with A. sachsi in
the Rio Negro and with A. balaenops in the Peruvian Am-
azon. A. sachsi is known to occur with A. devenanzii, both
in great numbers, and A. sachsi is expected to be found with
A. balaenops. The broader distributions of the phylogenet-
ically older species of A. clarkae and A. sachsi do not suggest
simple hypotheses on their speciation pattern and biogeo-
graphic history.

KEY TO SPECIES OF ADONTOSTERNARCHUS

la. Tail short (character not applicable to damaged or re-
generated individuals), head length contained less than
two times in caudal peduncle length; body depth below
origin of dorsal thong greater than or equal to least dis-
tance between eye and pectoral fin base; anal and pec-
toral fin membranes dark, or, a pale stripe on dorsal
midline 2

4 Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus

Figure 3. Head skeleton and pectoral girdle of Adontosternarchus devenanzii sp. n. 120.0 mm TL, MBUCV-V-4772. Lateral view of right
side. AN angular, BQ branchiostegal rays, CLT cleithrum, CU quadrate, DN dentary, ET ethmoid (=mesethmoid), ESFOT sphenotic, FR
frontal, HIO hyomandibular, I infraorbitals, IOP interopercular, MES mesopterygoid, MET metapterygoid, MIO intermuscular bones, MX
maxillary, OP opercular, ORS orbitosphenoid, PA parietal, PAS parasphenoid, POP preopercular, PMX premaxillary. PTM posttemporal,
RAR retroarticular, SCL supracleithrum, SIM symplectic, SOC supraoccipital, SOP subopercular, STC supratemporal sensory canal, VO
vomer.

lb. Tail long, head length usually contained more than two
times in caudal peduncle length; body depth below origin
of dorsal thong less than or equal to least distance be-
tween eye and pectoral hn base; anal and pectoral fin
membranes hyaline and no pale stripe on dorsal midline

3

2a. Interradial membranes of anal and pectoral fins darkly
pigmented with black or brown melanophores; no pale
stripe on dorsal midline; chin bulbous, often projecting
beyond snout; head profile nearly straight (Fig. 1A)

4. balaenops (Cope), Amazon Basin

2b. Interradial membranes of anal and pectoral fins hyaline;
a pale (yellow in life) stripe present on dorsal midline,
from snout (chin in most specimens) to near origin of
dorsal thong; chin rounded and little projected; head

profile rounded (Fig. IB)

A. devenanzii new species, Orinoco Basin

3a. Back and sides mottled with brown spots; anal rays ( 1 35?)
143-163 (Table 2); total pectoral rays 12-15 (Table 1);
snout length usually greater than interorbital distance

(snout 1.04-1.28 times interorbital width); body deeper,
maximum depth contained 3.6 to 5.1 times in distance

from snout to origin of dorsal thong (Fig. 9)

4. clarkae new species, Amazon Basin

3b. Back and sides nearly uniform brown (scale margins
have denser concentration of melanophores); anal rays
153-185 (Table 2); total pectoral rays 14-17 (Table 1 );
snout length usually less than interorbital distance (snout
0.80-1.04 times interorbital width); body shallower,
maximum body depth contained 4. 5-5. 8 times in dis-
tance from snout to origin of dorsal thong (Fig. 9) ... .
A. sachsi (Peters), Orinoco and Amazon Basins

Adontosternarchus balaenops (Cope, 1878)

Figures 1 A, 7-13

Sternarchus balaenops Cope, 1878:682 (original description,
single specimens). Eigenmann and Eigenmann, 1891:62
(listed).

Sternarchella balaenops. Eigenmann and Ward, 1905:164

Contributions in Science, Number 358

Mago-Leceia, Lundberg, and Baskin: Adontosternarchus 5

EHL

CHL

HHLD

Figure 4. Lateral view of the right lower hyoid apparatus of Adontosternarchus devenanzii sp. n. BQ branchiostegal rays, CHL ceratohyal,
EHL epihyal, HHLD dorsal hypohyal, HHLV ventral hypohyal, UH urohyal, IH interhyal.

(new combination, listed). Eigenmann, 1910:448 (listed).
Ellis, 1913:152 (copy of original description, bibliography).
Fowler, 1915: second page (characters). Fowler, 1 943: 121,
fig. 68 (profile of type, bibliography). Fowler, 1945:184,
fig. 68 (reprint of Fowler, 1943). Fowler, 1951:428 (bib-
liography).

Adontosternarchus sachsi. Ellis, 1913:156, pi. xxii, fig. 3 (in
part, Bolivia, San Joaquin, Rio Machupo). Eigenmann and
Allen, 1942:326 (in part, Peru, Iquitos). Fowler, 1939:278
(characters of single specimen, Peru, near Contamana, Rio
Ucayali). Fowler, 1951:423, fig. 465 (copied from Ellis,
1913).

Adontosternarchus balaenops. Eigenmann and Allen, 1 942:
327 (new combination, bibliography).

MATERIAL EXAMINED. Holotype of Sternarchus ba-
laenops: ANSP 21462, ca. 165 mm; Peru, Loreto State, Pe-
bas, Amazon River.

PERU: ANSP 83968, TL 138 (tail broken), Loreto State,
Ucayali River near Contamana.

BRAZIL: MBUCV-V- 1 1 522, 2, LEA 173.8 mm (other
length measurements not recorded due to damage); Ama-
zonas State, Lago Janauaca. MBUCV-V- 132 19, 1, TL 168.9
mm, LEA 1 50.0 mm (both measurements below normal due
to damage and regeneration), and MBUCV-V- 1 3220, 2, TL
190-232 mm; Amazonas State, Rio Solimoes, Ilha Mar-
chanteria, Lago Camaleao, 25 km SE of Manaus, Maria Ger-
cilia Mota. MCZ 9338, 1, TL 182 mm (measurement below
normal due to damage and regeneration); Amazonas State,
Rio Solimoes, Manacapuru. MZUSP 6896, TL 146.8 mm,
LEA 138.6 mm (both measurements below normal due to
damage and regeneration); Amazonas State, Rio Madeira,
25 km below Nova Olinda. MZUSP 24954, 3, TL 168.1-
1 88.9 mm, LEA 1 54.2-1 67.8 mm (measurements below nor-
mal due to damage and regeneration); Amazonas State, Rio
Solimoes, Lago Janauaca and vicinity. USNM 261385, 2,

Table 1. Frequency distributions of pectoral fin ray and caudal fin ray counts in Adontosternarchus.

Pectoral fin rays
(one fin counted per fish)

Caudal fin rays

12

13

14

15

16

17

18

10 11

12

13

14

15

16

17

18

19

20

21

22

balaenops

2

5

2

1

1

2

3

1

1

devenanzii

13

1 1

6

1

4

5

9

7

6

1

2

clarkae

5

7

6

3

1

2

1

6

4

sachsi

! 1

19

12

1

1

7

9

6

10

2

1

1

6 Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus

TL 151-163 mm, and USNM 229407, 1, TL 148.8 mm,
LEA 1 36.4 mm (measurements below normal due to damage
and regeneration); Amazonas State, Lago do Janauari, Lago
Terra Preta.

BOLIVIA; FMNH 54568 (formerly CM 3199), 5 (origi-
nally 6), TL 122.6-163.7 mm, LEA 1 13.5-139.5 mm (both
measurements below normal due to damage and regenera-
tion); Beni State, San Joaquin, Machupo. UMMZ 204883,
9, TL 106-124 mm; Beni State, Rio Baures, 2 km above
mouth.

DIAGNOSIS. Tail and caudal peduncle short, head length
contained less than two times in peduncle; dorsal thong does
not reach end of anal fin; body deep, maximum depth 222-
385 thousandths of length to origin of dorsal thong (see Fig.
9); depth below origin of dorsal thong exceeds or equals least
distance between eye and pectoral base; head angular, its
dorsal profile sloping in nearly straight or slightly concave
line to snout; chin projecting (Figs. 1A, 8), interorbital dis-
tance does not reach from eye to tip of chin; distance to anal
fin origin 172-233 thousandths of length to origin of dorsal
thong; 143-179 anal rays (Table 2); 1 5-1 8 pectoral rays (Ta-
ble 1 ); 1 3-2 1 caudal rays (Table 1 ); back and sides irregularly
marked with spots and blotches; no pale dorsal midline stripe;
pectoral and anal fin interradial membranes dark brown or
black.

DESCRIPTION. Measurements in thousandths of refer-
ence dimension. Body depth about 162-196 of LEA, 225-
315 of length to origin of dorsal thong, 190-228 of anal fin
base 1108-1521 of head length; depth at nape 152-214 of
length to origin of dorsal thong, 767-1032 of head length;
dorsal profile of body gently convex, more so behind the
head; preanal fin distance 137-167 of LEA, 156-195 of anal
fin base, 171-242 of length to origin of dorsal thong (Fig.
1 1 ); caudal peduncle 1 1 4-1 6 I of LEA, 1 33-1 84 of anal fin
base.

Head somewhat compressed, its length 1 7 1-223 of length
to origin of dorsal thong; distance from snout tip to rictus
673-1056 of snout length; chin rounded; end of snout bluntly
pointed; snout length 254-295 of head length, 818-1 1 1 I of
interorbital distance; both jaws edentulous in adults.

Eye diameter 75-115 of head length, 272-406 of snout
length, 269-377 of interorbital distance; interorbita! distance
254-323 of head length; branchial opening 197-345 of depth
at nape, 169-293 of head length; distance from tip of snout
to vent 82-95 of LEA, vent shifts relatively anteriad with
growth (Fig. 12).

Figure 5. Upper pharyngeal tooth plate (PFS) and associated bones
of Adontosternarchus devenanzii sp. n. EPI 1-5 epibranchials, I 2-
4 infrapharyngobranchials. EPI 5 and I 4 are cartilaginous. 1 1 is
absent.

Anal fin base about 857-913 of LEA; length of pectoral
fin 725-965 of head length.

53-60 vertebrae to base of last anal fin ray.

Background color in alcohol tan to brown; sides and back
mottled with brownish-black chromatophores; spots and large

Table 2. Frequency distribution of anal fin rays in Adontosternarchus.

Anal fin rays (grouped by twos)

135 137 139 141 143 145 147

149

151

153 155

157

159

161

163

165 167

169 171

173 175 177 179 181 183 185

balaenops

1 1

2

3

2

1

2

1

1

2

1

devenanzii

2

2

2

8 5

6

4

5

2

1

clarkae

1 I 3

2

1

6 1

2

2

2

sachsi

1 1

1

2

4

3 4

4

3 5 3 7 11

Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus 7

SCL+PTM

CLT

Figure 6. Medial view of the left pectoral girdle of Adontotosternarchus devenanzii sp. n. CLT cleithrum, COR coracoid, ESC scapula, PCL
postcleithra, PTM posttemporal, RP pectoral radials, SCL supracleithrum.

dense blotches scattered irregularly on sides and dorsum;
dorsal midline without a pale stripe; dorsal thong with hya-
line ground color but otherwise colored as the back; lateral
line sensory canal evident as a thin pale broken line on sides;
lower sides with scattered superficial spots underlain with
numerous dark, ventroposteriorly oblique lines formed by
deep chromatophores and spaces between anal bases; anal

fin membrane mostly black but few or no chromatophores
over rays; caudal fin (all regenerated in material examined)
mostly hyaline or with a light peppering of chromatophores
at the base; anal and pectoral fin membranes mostly black
except over the rays; top and upper sides of head pigmented
as the body, tip of snout with a pale area; chin dusky or pale;
lateral margin of upper lip pale; sides and under surface of

Figure 7. Lateral view of Adontosternarchus balaenops (CM 3 1 99, 140 mm); reproduced from Ellis (1913, plate XXII, fig. 3) with permission
of the Carnegie Museum.

8 Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus

o □

Figure 8. Adontosternarchus balaenops (Cope), 1 74.0 mm TL, Lago
Janauaca, Rio Solimoes, Amazonas, Brazil. A, entire fish; B, close
up of head.

head paler, with variable amount of scattered chromato-
phores; tube of anterior naris immaculate.

DISTRIBUTION. Adontosternarchus balaenops is thus far
known from the lowlands (< ca. 200 m elevation) of the
Amazon River Basin of Brazil, Peru, and Bolivia (Fig. 13).

REMARKS. The original description of A. balaenops was
based on a single specimen that had become severely dis-
torted and damaged through desiccation. Ellis, following Ei-
genmann and Ward (1905), assigned the species to Sternar-

36 48 60 72 84 96 108 120

DISTANCE TO ORIGIN OF DORSAL THONG IN MM

Figure 9. Greatest body depth versus distance to origin of dorsal
thong in Adontosternarchus; closed squares, balaenops; open squares,
clarkae; closed circles, sachsi; open circles, devenanzii.

f •

° 8

60 72 84 96 108 120

DISTANCE TO ORIGIN OF DORSALTHONG IN MM

Figure 10. Caudal peduncle length versus distance to origin of
dorsal thong in Adontosternarchus; closed squares, balaenops; open
squares, clarkae; closed circles, sachsi; open circles, devenanzii.

5

DISTANCE TO ORIGIN OF ANAL FIN IN MM

Figure 11. Length to origin of dorsal thong versus distance to origin
of anal fin in Adontosternarchus; closed squares, balaenops; open
squares, clarkae; closed circles, sachsi; open circles, devenanzii.

4° * S

■ «o °»

■ ° i> ? .

.. *.
’ oo o\*

t ' i i i i i i i ! i i i i r

36 48 60 72 84 96 108 120

DISTANCE TO ORIGIN OF DORSAL THONG IN MM

Figure 12. Snout to vent length versus distance to origin of dorsal
thong in Adontosternarchus; closed squares, balaenops; open squares,
clarkae; closed circles, sachsi; open circles, devenanzii.

chella, a genus defined in part by the presence of small teeth
in both jaws. The holotype, however, shows neither gnathal
dentition, nor the nearly straight dorsal profile of Sternar-
chella (Ellis, 1913:151, fig. 14), but does exhibit features of
the distinctive snout and jaws of Adontosternarchus (Fig. 1 ).
Without explicit justification Eigenmann and Alien (1942)
transferred balaenops to Adontosternarchus. One feature of

Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus 9

90

00

70

^ 60

50

Figure 13. Geographic distribution of Adonlosternarchus balaenops (B) and A. devenanzii ( D); circled symbols = type localities; some symbols
represent more than one collection locality or lot of specimens (see especially text section on distribution under A. devenanzii and Fig. 15).

the holotype that early concerned us as not being a character
of Adontosternarchus is its concave, rather than bulbous chin.
This, however, is clearly a result of desiccation at some point
in the specimen’s history; we find exactly the same concave
chin shape with projecting lower lip in purposely dried spec-
imens of Adontosternarchus. We fully agree with Eigenmann
and Allen's generic placement of balaenops.

Cope’s description of the type of balaenops offers little help
in determining its relationship to the material examined by
us. He wrote (1878:682),

“Profile oblique, with a depression between the orbits;
snout short and much narrowed; lower jaw large, pro-
jecting beyond the upper both anteriorly and laterally,
enclosing the latter somewhat as in a whalebone whale.
The fissure of the mouth is short, only reaching the ver-
tical line from the anterior nostril. Eyes small, without
free border, much nearer the snout than the gill-opening,
one twelfth the length of the head, which latter enters

the length without the caudal fin, 8.5 times. The depth
at the base of the dorsal thong is equal to the length of
the head. Anal radii 171. Scales very large, in only nine
longitudinal rows at the base of the dorsal thong. Color
olivaceous, with a pale dorsal band which reaches the
dorsal thong, and a pale narrow band on each side near
the dorsal band. Length 165 mm.; length to origin of
anal 20 mm.; length to base of dorsal thong 96 mm.

This species resembles remotely the S. schotti of Stein-
dachner, but differs from it and from all the other species
in the much enlarged mandible and large scales.”

Most of these features suggest only Adontosternarchus in gen-
eral, with some added specimen damage. Cope’s count of
1 7 1 anal radtals (the fin is missing) may be accurate but it
falls within the anal fin ray count range of the two of the
other three species recognized herein. (Our count of 1 56 ra-
dials and length measurement of 143 mm indicates that a
part of the tail is now missing.) Cope’s color notes could be

10 Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus

positively misleading since the pale dorsal bands are created
by exposed connective tissue of myosepta; these are not part
of the integumentary pigmentation and thus bear no simi-
larity to the pale dorsal band of A. devenanzii. In fact, given
the poor state of the type nothing can be said of its skin or
fin pigmentation. Contrary to Cope's remark, the depth of
the body at the base of the dorsal filament does not nearly
equal the head length unless he intended to exclude snout
and eye. This depth, however, exceeds the least distance
between the eye and pectoral base, a character which we have
found to be diagnostic of some species. Furthermore, the
maximum body depth is contained a little less than 4 V2 times
in the length to the origin of the dorsal thong (Fig. 9). These
two expressions of a relatively deep body characterize two
otherwise distinguishable species of Adontosternarchus: the
Orinocan A. devenanzii, and the central Amazonian species
to which we here apply the name A. balaenops. Our decision
is based on the depth measurements and known geographic
distribution of the two species. Other possible diagnostic
features are not preserved in the type. Ours is a conservative
course that avoids coming a likely superfluous binomen. We
do recognize that, if the two species are someday found sym-
patric and no other diagnostics are discovered, balaenops
might become a nomen dubium.

Adontosternarchus devenanzii new species

“De Venanzi’s knifefish”

Figures IB, 2-7, 9-15

Adontosternarchus sachsi. Mago-Leccia, 1967:257, fig. 10
(Venezuela). Mago-Leccia, 1970:77 (listed). Adontoster-
narchus sp. Lopez, Lundberg, and Marsh, 1984:333 (Ven-
ezuela, Rio Orinoco Delta).

HOLOTYPE. MBUCV-V-75 1 3, 1, TL 133.4 mm, LEA
1 13.0 mm (see Table 3); Venezuela, Cano Caujarito, tribu-
tary of Rio Portuguesa, 3 km above La Union, Guarico State,
23 Aug. 1974, J.N. Baskin, J.O. Silva, and L. Aguana.

PARATYPES. VENEUZUELA: MBUCV-V-4772, 20, TL
7 1 .2-159.8 mm; Cano Caracara tributary of Rio Meta, Apure
State, 10 March 1967, F. Mago-Leccia and J. Mosco (4 spec-
imens cleared and stained). MBUCV-V-5 1 39, 1, TL 106.8
mm; Cano Cocuiza, Tabirito bridge, near Caicara del Ori-
noco, Bolivar State, 27 Feb. 1 969, F. Mago-Leccia and party.
MBUCV-V-5984, 1, TL 1 16.5 mm; Esteros de Camaguan,
Guarico State, 6 Aug. 1971, A. Machado. MBUCV-V-75 16,
4, TL 1 1 1.0-135.9 mm; Boca Ruido lagoon, Rio Portuguesa
system, Guarico State, 8 Aug. 1971, F. Mago-Leccia.
MBUCV-V-9301, 5, TL 85.9-123.7 mm; Modulos de
Mantecal, Apure State, 1 June 1974, L. Aguana and A. Ma-
chado. MBUCV-V-12701, 4, TL 146.2-186.2 mm; Esteros
de Camaguan, Guarico State, 25 April 1980, F. Provenzano,
O. Castillo, and L. Aguana. The following material collected
by J.N. Baskin, J.G. Lundberg, and F. Mago-Leccia. MBUCV-
V-10385, 3, TL 154.6-162.5 mm; Rio Orinoco, Los Castil-
los, southside of channel. Delta Amacuro Territory, 16 Feb.
1978. MBUCV-V- 10486, 1, TL 137.7 mm; Rio Orinoco,
main channel North of Isla Tres Canos, at Cano Araguaito,

Table 3. Measurements in mm and thousandths of length to end of
anal fin base (LEA), and counts of meristic characters for holotypes
of Adontosternarchus devenanzii and A. clarkae.

Measurements

A. devenanzii
MBUCV-V-75 13

A. clarkae
MBUCV-V-12703

mm

mils LEA

mm

mils LEA

Total length

133.4

1181

172.0

1

536

LEA

1 13.0

-

1 12.0

-

Maximum body depth

16.6

147

17.4

155

Head depth at nape

12.9

1 14

13.4

120

Length to origin of anal fin

17.6

156

15.1

135

Caudal peduncle length

13.2

117

52.9

472

Tail length

20.4

181

55.0

491

Length to origin of dorsal

68.2

608

80.9

722

thong

Head length

16.0

142

12.7

113

Snout length

4.5

40

3.4

30

Eye diameter

1.7

15

1.4

13

Interorbital distance

4.8

42

3.9

35

Postorbita! length

10.4

92

8.5

76

Size of branchial aperture

3.1

27

2.2

20

Length to anus

7.6

67

6.9

62

Length of anal fin base

99.0

876

101.2

904

Pectoral fin length

12.5

1 1 1

10.8

96

Pectoral fin rays

ii.

13-14

ii, 10

Anal fin rays

153

154

Caudal fin rays

20

16

Delta Amacuro Territory, 19 Feb. 1978. MBUCV-V- 10497,
6, TL 115.1-144.5 mm; Rio Orinoco, upstream from Cano
Tres Canos, Delta Amacuro Territory, 19 Feb. 1978.
MBUCV-V- 10528, 1, TL 152.9 mm; Rio Orinoco, 2 km
downstream from Barancas, Delta Amacuro Territory, 17
Feb. 1978. MBUCV-V-10535, 3, TL 126.8-174.7 mm; Rio
Orinoco, Los Castillos, Delta Amacuro Territory, 16 Feb.
1978. MBUCV-V- 10580, 1, TL 125.6 mm; Rio Orinoco,
Tapatapa lagoon in Isla Tapatapa, Delta Amacuro Territory,
16 Feb. 1978. MBUCV-V- 10588. 2, TL 103.4-159.0 mm;
Rio Orinoco, Brazo Imataca, Delta Amacuro Territory, 22
Feb. 1978. MBUCV-V-10595, 3, TL 1 10.8-132.0 mm; Rio
Orinoco, main channel N of Isla Tres Canos at Cano Ara-
guaito, Delta Amacuro Territory, 19 Feb. 1978. USNM

Table 4. Frequency distributions of number of vertebrae to base of
last anal fin ray in Adontosternarchus.

57

58

59

60 61

62

63

64

balaenops

2

1

4

2

devenanzii

2 3

2

2

2

clarkae

2

2

4

1 1

sachsi

1

1 2

4

3

Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus 1 1

Figure 14. Adontosternarchus devenanzii sp. n., 154.6 mm TL, MBUCV-V-10385, paratype, Orinoco River, Los Castillos, Venezuela. A,
entire fish; B, close up of head.

264839, 68, LEA 84-1 17 mm; Rio Orinoco, main channel
S of Isla Portuguesa, Delta Amacuro Territory, 20 Feb. 1978.
LACM 43103-1, 75, LEA 90-147 mm; Rio Orinoco, main
channel N of Isla Fajardo, Delta Amacuro Territory, 14 Feb.
1978. FMNH 94909, 12, LEA 82-130 mm; Rio Orinoco,
main channel N of Isla Tres Canos, 19 Feb. 1978. CAS
54328, 24 LEA 82-124 mm; Rio Orinoco, main channel
near Cano Remolina, 25 Feb. 1978. LACM 43295-1, 27,
LEA 67-84 mm; Rio Orinoco, north shore at Isla Portuguesa
in Cano Anabata, Delta Amacuro Territory, 16 Nov. 1979.

COLOMBIA: ANSP 128203, 8, TL 88-130 mm, LEA 76-
1 1 1 mm; Meta State hacienda Mozambique, Laguna Mo-
zambique, N shore, 20 March 1971, J.E. Bohlke. ANSP
1 3 1836, 1 , TL 127 mm, LEA 1 14 mm, Meta State, Quebrada
Venturosa between La Balsa and Puerto Lopez, 4°05'N,
72°58'W, 21 March 1975, J.E. Bohlke. ANSP 138859, 1,TL
123 mm, LEA 109 mm; Meta State, Rio Metica, N bank of
river SW of Laguna Mozambique, 20 Feb. 1972, J.E. Bohlke.

OTHER MATERIAL. Additional material of A. deve-
nanzii comprising 89 lots and 4594 individuals was collected

12 Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus

Feb

DEVENANZII

Deep River

82

52

28

35 1103

851

11

12

111

57

65

1

98.8

(n=2438)

Shallow River

23

0.9

1978

Near Shore

3

2

1

1

0.3

(low

SACHSI

(n=652)

Deep River

67

73

111

19 120

162

1

1

30

36

6

10

16

100.0

water)

Shallow River
Near Shore

DEVENANZI I

Deep River

2

12

1

4

1

0.3

Nov

(n=2184)

Shallow River

6

4 2031

99

10

99,1

1979

Near Shore

3

3

2 1

5

0.6

(HIGH

Deep River

5

4

7

8 1.8

water)

SACHSI

Shallow River l

14

117

7

1014

100

18

5

37

96.9

(n=1359)

Near Shore

21

1

1.6

Figure 15. Longitudinal and habitat distribution of Adontosternarchus sachsi and A. devenanzu in the lower Orinoco River in 1978 and
1979. Values in the table are numbers of specimens taken in the indicated habitat (rows) at the indicated site (columns). (See text section on
distribution under A. devenanzii for discussion.)

in 1978 and 1979 in the Orinoco River delta region, Vene-
zuela, by J.N. Baskin, J.G. Lundberg, and F. Mago-Leccia
from R/V EASTWARD, then of Duke University. This ma-
terial is deposited in the following institutions: USNM,
LACM, FMNH, UMMZ, ANSP, AMNH, and MCZ.

DIAGNOSIS. Tail and caudal peduncle short, head length
contained in peduncle less than twice; dorsal thong usually
does not reach end of anal fin (sometimes extends slightly
beyond); body deep, maximum body depth 208-286 thou-
sandths of length to origin of dorsal thong (Fig. 9); depth
below origin of dorsal thong exceeds or equals least distance
between eye and pectoral fin base; head chubby and rounded,
its dorsal profile distinctly convex; chin not markedly pro-
jecting (Figs. IB, 14), interorbital distance reaches from eye
to tip of chin or beyond; distance to origin of anal fin 1 70—
256 thousandths of length to origin of dorsal thong; 147-173
anal rays (Table 2); 14-16 pectoral rays (Table 1); 15-22
caudal rays (Table 1); a pale (yellow in life) stripe from chin
tip and snout to near origin of dorsal thong (this stripe is
occasionally obscured by dark spots or is very rarely obso-
lescent); pectoral and anal fins hyaline.

DESCRIPTION. Measurements in thousandths of refer-
ence dimension. Body depth 131-179 of LEA, 208-283 of
length to origin of dorsal thong, 1 38-206 of anal fin base and
104-160 of head length; depth at nape 85-124 of LEA, 1 37—
206 of length to origin of dorsal thong, 90-140 of anal fin
base, 765-1020 of head length; dorsal profile of body gently
convex, more so behind the head (Fig. 14); preana! fin dis-
tance 119-159 of LEA, 195-256 of distance to origin of
dorsal thong (Fig. 1 1), 126-183 of anal fin base; caudal pe-
duncle length 79-204 of LEA, 131-313 of distance to origin

of dorsal thong (Fig. 10), 90-228 of anal fin base and 70-
168 of total length; tail length 121-236 of LEA, 194-386 of
distance to origin of dorsal thong, 133-267 of anal fin base,
108-191 of total length; length to origin of dorsal thong 583-
716 of LEA.

Head rounded; head length 107-138 of LEA, 164-209 of
distance to origin of dorsal thong, 1 13-158 of anal fin base
and 578-1435 of caudal peduncle; dorsal profile of head
convex, smoothly continuous with contour of back (Fig. 14),
except in individuals ca. 100 mm TL in which there is often
a shallow concavity at the nape; distance from snout tip to
rictus 600-953 of snout length; chin rounded (Fig. IB); end
of snout bluntly pointed; snout length 260-298 of head length,
771-1040 of interorbital distance; both jaws edentulous in
adults, but carrying conical teeth in juveniles up to ca. 26
mm TL.

Eye diameter 86-142 of head length, 317-500 of snout
length, 265-500 of interorbital distance; interorbital distance
250-36 1 of head length; branchial opening 153-29 1 of depth
at nape, 1 52-244 of head length; distance from tip of snout
to vent 58-99 of LEA, vent shifts relatively anteriad with
growth (Fig. 12).

Anal fin base 802-976 of LEA; length of pectoral fin 829-
969 of head length.

56-64 vertebrae to base of last anal fin ray.

Background color in alcohol pale tan to yellowish-white;
sides and back densely mottled with brownish-black chro-
matophores; spots and large dense blotches scattered irreg-
ularly on sides and dorsum; dorsal midline with a pale stripe
from chin and snout to near origin of dorsal thong, irregularly
obscured by dark spots or rarely obsolescent; dorsal thong

Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus 13

with a hyaline ground color but superficially spotted; lateral
fine sensory canal evident as a thin, pale broken line on sides;
lower sides with scattered superficial spots underlain with
numerous dark, ventroposteriorly oblique lines formed by
deep chromatophores and spaces between anal fin bases; anal
fin membrane hyaline except usually a few chromatophores
present over rays; caudal fin usually with scattered chro-
matophores, particularly on its base; pectorals hyaline; top
and upper sides of head pigmented as the body, the pale
stripe of the dorsal midline extending onto tips of snout and
chin where it is surrounded by a dark U-shaped band; sides
and under surface of head paler, with scattered chromato-
phores; lateral margin of upper lip pigmented; tube of an-
terior naris always bears a few chromatophores.

Color in life; background color dark brownish, mottled;
head yellow ocher and dark brown; dorsal midline with a
yellow stripe; fins all hyaline, except the caudal which is
pigmented with a darker area on its base; opercles darker
than surrounding areas; snout with dark areas on each side
of the midline stripe; chin with a dark U-shaped band.

ETYMOLOGY. The name devenanzii is for Dr. Francisco
De Venanzi, former Rector of the Universidad Central de
Venezuela, Caracas, who encouraged the first author to study
fishes.

DISTRIBUTION. This new species is quite common
throughout the Venezuelan and Colombian Low Llanos, the
main course of the Rio Orinoco and its large tributaries (Fig.
13).

Collections made by us in the Orinoco Delta region in

1978 and 1979 reveal that Adontosternarchus are more abun-
dant than prior sampling suggested. In our survey work col-
lections were made with trawls from R/V EASTWARD in
deep river channels (10-80 m) and from smaller craft in
shallower areas (ca. 10 m), as well as with conventional col-
lecting gear in near shore habitats. The 1978 expedition was
in February during the middle of the low water (dry) season;
the 1979 expedition was in November near the end of the
high water (wet) season. Large numbers of A. devenanzii and
A. sachsi were collected in both years and their distributions
along the transects were similar (Fig. 15).

Between 46% and 96% of the total sample of each species
in each year were collected between channel markers 120
n,mi and 140 n,mi, the 20 n,mi stretch of river just below
the head of the delta (channel markers and lights of the ship-
ping lanes are labelled with distances from the sea buoy which
is located about 30 n,mi off Boca Grande). A. devenanzii was
not taken downstream from the 60 n,mi marker or above
the 201 n,mi marker, but A. sachsi was found further down-
stream to the 42 n,mi marker (just 7 n,mi above the transition
to brackish water) and further upstream at 24 1 n,mi. Despite
the between-year similarity in longitudinal distribution, both
species appear to shift habitat between low and high water
seasons. In 1978 (low water) 88.7% of the A. devenanzii and
100% of A. sachsi were collected in bottom trawls in mid-
river channels usually much greater than 10 m in depth. In

1979 only 11.5% of devenanzii and 28.6% of sachsi were
collected in deep channels even though trawling effort in this
habitat was greater in that year (151 vs. 186 deep channel

trawls). The largest numbers of Adontosternarchus collected
in 1 979 (57.7% of devenanzii and 62.9% of sachsi ) were taken
with a small trawl pulled by a dugout canoe in lagoons and
shallow places of the river between about 30 and 110 m from
shore (Lopez, Lundberg, and Marsh, 1984). In both years
these fishes were rarely captured in near shore habitats. Over-
all then, it appears that Adontosternarchus are bottom-ori-
ented fishes of large rivers and lagoons and that they make
marked seasonal movements between the deep channels and
the shallow channels and lagoons. The biological significance
of this movement remains to be investigated.

Adontosternachus clarkae new species

“Clark’s knifefish”

Figures 1C, 9-12, 16-17

HOLOTYPE. MBUCV-V- 12703, TL 172.0 mm, LEA
1 1 2.0 mm (see also Table 3); Venezuela, Raudal (Rapids) de
Mavahate, Rio Negro, near San Carlos de Rio Negro, Ama-
zonas Territory, 6 May 1981, Edgar Armas.

PARATYPES. VENEZUELA: MAC-PAY-0369, 3, TL
172-186 mm, LEA 1 10-1 19 mm; paratopotypes taken with
the type specimen. MBUCV-V-1 1218, 2, TL 158.6-172.5
mm; rapids downstream from the mouth of the Casiquiare
in the Rio Negro, about 10 km N of San Carlos de Rio Negro,
Amazonas Territory, 2 May 1 978, Kate Clark and Raimundo
Videra. MBUCV-V- 1 1 293, 1 , TL 1 62.4 mm; San Carlos de
Rio Negro, Amazonas Territory, Nov. 1980, Kate Clark.
MBUCV-V- 1 1 936, 2, TL 168.1-168.5 mm; paratopotypes
(one specimen stained).

COLOMBIA: FMNH 94263, 5, TL 73.1-94.8 mm, LEA
68.1-77.4 mm; Amazonas State, Amazon River above Le-
ticia, Nov. 1973, .1. Thomerson.

PERU: MZUSP uncatalogued, 2, TL 88.3-133.8 mm, LEA
72.5-109.0 mm; Ucayali State, Rio Ucayali, Masisea, H.
Ortega, Oct. 1975.

ECUADOR: FMNH 94264, 1, TL 105.3 mm, LEA 75.5
mm; Rio Tiputini at confluence with Rio Napo, 0°48.9'S,
75°32.5'W, Dec. 1981, D.J. Stewart.

BRAZIL: MCZ 46877, 14 1 , TL 85.0-104.2 mm, LEA 65.3-
80.5 mm; Terr. Roraima, Rio Negro near confluence with
Rio Branco and lower 30 miles of Rio Branco, 1°24'S,
6 1°27'W, March-April 1967, Alpha Helix. USNM 266551,
3. TL 95-127 mm; same data as preceding lot. MCZ 46872,
41, TL 60.4-90.9 mm, Rio Negro at 1°24'S, 61°27'W, 2 miles
below confluence with Rio Branco, 27-28 March 1967, Al-
pha Helix. LACM 43645- 1 , 3, TL 74. 1-94. 1 mm; same data
as preceding lot.

DIAGNOSIS. Tail and caudal peduncle long, head length
contained in peduncle twice or more; dorsal thong reaches
beyond end ofanal fin; body depth moderate relative to other
species of the genus, maximum body depth 196-278 thou-
sandths length to origin of dorsal thong (Fig. 9); depth below
origin of dorsal thong less than or equal to least distance
between eye and pectoral base; head angular, its dorsal profile
straight or slightly convex; chin projecting (Figs. 1C, 16),
interorbital distance reaches from eye to chin tip; distance
to origin of anal fin 167-256 thousandths of length to origin

14 Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus

Figure 16. Adontosternarchus clarkae sp. n., 172.5 mm TL, MBUCV-V-1 1218, paratype, Rio Negro, Amazonas Territory, Venezuela. A,
entire fish; B, close up of head.

of dorsal thong; (135 partly regenerated?) 144-164 anal fin
rays (Table 2); 12-15 pectoral fin rays (Table 1 ); 10-16 caudal
fin rays (Table 1 ); back and sides mottled with light and dark
brown, oddly shaped spots; no pale dorsal midline stripe;
pectoral and anal fins hyaline.

DESCRIPTION. Measurements in thousandths of refer-
ence dimension. Body depth 138-170 of LEA, 195-278 of
length to origin of dorsal thong, 154-199 of anal fin base.

1 149-1458 of head length; depth at nape 92-127 of LEA,

1 26-183 of length to origin of dorsal thong, 105-149 of anal
fin base, 730-1 1 18 of head length; dorsal profile of body
gently convex more so behind the head (Fig. 16); preanal
distance 1 15-154 of LEA, 166-259 of distance to origin of
dorsal thong (Fig. 1 1), 133-175 of anal fin base; caudal pe-
duncle length 258-495 of LEA, 379-679 of distance to origin
of dorsal thong (Fig. 10), 293-549 of anal fin base, 194-318

Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus 15

Figure 17. Geographic distribution of Adontosternarchus clarkae(C) and A. sachsi (S); circled letters = type localities; some symbols represent
more than one collection locality or lot of specimens (see especially text section on distribution under A. devenanzii and Fig. 1 5 for additional
information on A. sachsi).

of total length; tail length 310-565 of LEA, 454-765 of dis-
tance to origin of dorsal thong, 351-633 of anal fin base,
243-361 of total length; distance to origin of dorsal thong
595-756 of LEA.

Head rounded (head length 104-140 of LEA, 139-228 of
length to origin of dorsal thong, 117-158 of anal fin base,
213-495 of caudal peduncle); dorsal profile of head slightly
convex, smoothly continuous with contour of back (Fig. 1C);
distance from snout tip to rictus 543-1000 of snout length;
chin bulbous; end of snout bluntly pointed; snout length 253-
303 of head length, 775-1000 interorbital distance; both
jaws edentulous in adults.

Eye diameter 85-130 of head length, 294-500 of snout
length, 267-458 of mterorbital distance; interorbital distance
270-345 of head length; branchial opening 1 28-268 of depth
at nape, 137-242 of head length; distance from tip of snout
to vent 54-90 of LEA, vent shifts relatively anteriad with
growth (Fig. 12).

Anal fin base 851-919 of LEA; length of pectoral fin 833-
1021 of head length.

57-61 vertebrae to base of last anal fin ray.

Background color in alcohol pale tan to yellowish-white;
sides and back densely mottled with irregularly scattered,
brownish-black blotches; dorsal midline without a pale stripe;

dorsal thong with a hyaline ground color but superficially
with dark brown blotches; lateral line sensory canal evident
as a thin pale broken line on sides; anal fin membrane hyaline
except for a few chromatophores present over rays; caudal
fin pigmented with small spots and chromatophores more
concentrated over its base; pectoral fins hyaline; top and
upper sides of head with blotches as the body; lower sides
and undersurface of head with scattered blotches and dots;
tip of snout, margin of upper lip, tip of chin, and tube of
anterior naris always immaculate.

ETYMOLOGY. The name clarkae is for Ms. Kate Clark,
collector of this new species from the Rio Negro, Venezuela.

DISTRIBUTION. Adontosternarchus clarkae is distrib-
uted in the upper parts of the Amazon River Basin of Brazil,
Venezuela, Colombia, Ecuador, and Peru (Fig. 17). There
are no records of it from the lower Amazon. The Venezuelan
specimens were collected in black waters near rapids.

Adontosternarchus sachsi (Peters)

“Sachs’ knifefish”

Figures ID, 9-12, 15, 17-19

Sternarchus sachsi Peters, 1877:473 (original description, type
locality: San Fernando de Apure, Venezuela). Sachs, 1879:

16 Contributions in Science, Number 358

Mago-Leecia, Lundberg, and Baskin: Adontosternarchus

Figure 18. Adontosternarchus sachsi (Peters) (exact origin and disposition of specimen unknown) reproduced from Sachs (1879:279).

153, 367, fig. on p. 279 (Apure). Eigenmann and Eigen-

mann, 1891:62 (listed). Rohl, 1942:377, fig. 189 (copied
from Sachs).

Sternarchogiton sachsi. Eigenmann and Ward, 1905:165 (new
combination). Ihering, 1 907:275 (listed). Eigenmann, 1910:
448 (listed).

Adontosternarchus sachsi. Ellis, 1913:156 (in part). Fowler,
1939:278 (Peru, Contamana). Schultz, 1949:74 (listed,
characters). Fowler, 1943:124 (listed). Fowler, 1945:185
(reprint of Fowler, 1943). Fowler, 1951:423 (listed).

MATERIAL EXAMINED. VENEZUELA: Photograph
of holotype of Sternarchus sachsi; ZMB No. 10044; Vene-
zuela, Apure State, San Fernando de Apure. MBUCV-V-
11292, 1, TL 165.6 mm; Venezuela, Rio Negro, Amazonas
Territory, near San Carlos de Rio Negro. MBUCV-V-4643,
1, TL 120.8 mm; Rio Orinoco, Quiritare, Amazonas Ter-
ritory. MBUCV-V- 10377, 3, TL 152.8-200.5 mm; Rio Or-
inoco, Brazo Imataca, Delta Amacuro Territory. MBUCV-
V- 1 04 1 7, 2, TL 1 90.8-203.0 mm; Rio Orinoco, old shipping
channel, S of Isla Portuguesa, Delta Amacuro Territory.
MBUCV-V- 10431, 3, TL 211.1 —222.9 mm; Rio Orinoco,
near Isla Iguana, Delta Amacuro Territory. MBUCV-V-
10441, 2, TL 193.4-202.6 mm, Rio Orinoco, along S shore
by Cairo Guine, Delta Amacuro Territory. MBUCV-V- 1 0445,
1, TL 186.4 mm; Venezuela, Rio Orinoco, along S shore by
Cairo Guine, Delta Amacuro Territory. MBUCV-V-10470,

I, TL 169.9 nrnr; Rio Orinoco, main channel E of Isla Por-
tuguesa, Delta Amacuro Territory. MBUCV-V- 10495, 3, TL
147.8-180.7 mm; Rio Orinoco, Cairo Tres Canos, Delta
Amacuro Territory. MBUCV-V- 10506, 1, TL 200.0 nrnr;
Rio Orinoco, near Los Castillos, Delta Amacuro Territory.
MBUCV-V- 105 54, 1 3, TL 145.5-32 1 .8 mm; Venezuela, Rio
Orinoco, Isla Veradero, Delta Amacuro Territory (4 speci-
mens stained). LACM 43295-2, 29, LEA 57-85 nrnr; Rio
Orinoco, north shore at Isla Portuguesa in Cairo Anabata,
Delta Amacuro Territory.

BRAZIL: MZUSP 24925, 2, TL 138.2-160.2 nrnr, LEA
1 29.4-14 1 .6 nrm Rio Solinroes, Lago Janauaca and vicinity.
FMNH 54569 and 1 5 1 87, 1 5 1 88, 1 5 1 89, 1 5 190, 1 5 1 9 1 (for-
merly CM 3200), 55, TL 93.4-1 16.6 mm, LEA 83.2-108.3
mm; Para State, Santarenr.

OTHER MATERIAL. Additional material of A. sachsi
comprising 89 lots and 201 1 individuals was taken in 1978
and 1979 in the Orinoco River Delta region, Venezuela, by

J. N. Baskin, J.G. Lundberg, and F. Mago-Leccia from R/V
EASTWARD, then of Duke University. This material is de-

posited in the following institutions: USNM, LACM, FMNH,
UMMZ, CAS, ANSP, AMNH, and MCZ.

DIAGNOSIS. Tail and caudal peduncle long, head con-
tained in peduncle twice or more; dorsal thong reaches be-
yond end of anal fin; body shallow, maximum body depth
172-222 thousandths of distance to origin of dorsal thong
(Fig. 9); depth below origin of dorsal thong less than or equal
to least distance between eye and pectoral base; head angular,
its dorsal profile variable but not strongly convex; chin pro-
jecting (Fig. 1 D, 19), interorbital distance not reaching from
eye to chin tip; distance to origin of anal fin 137-213 thou-
sandths of distance to origin of dorsal thong; 154-185 anal
rays (Table 2); 14-17 pectoral rays (Table 1); 14-22 caudal
rays (Table 1); back and sides nearly uniform brown, except
for dark margins of some scales; no pale midline stripe; pec-
toral and anal fins hyaline.

DESCRIPTION. Measurements in thousandths of refer-
ence dimension. Body depth 116-143 of LEA, 174-218 of
distance to origin of dorsal thong, 126-174 of anal fin base,
953-1548 of head length; depth at nape 79-106 of LEA,
1 1 9-1 63 of distance to origin of dorsal thong, 85-129 of anal
fin base, 733-1087 of head length; dorsal profile of body
scarcely convex to straight (Fig. 1 9); preanal distance 86-146
of LEA, 137-214 of distance to origin of dorsal thong (Fig.
11), 93-169 of anal fin base; caudal peduncle length 219-
366 of LEA, 327-524 of distance to origin of dorsal thong
(Fig. 10), 236-404 of anal fin base, 170-257 of total length;
tail length 287-426 of LEA, 41 1-628 of distance to origin
of dorsal thong, 306-468 of anal fin base, 217-299 of total
length; distance to origin of dorsal thong 621-710 of LEA.

Head slightly compressed (head length 92-135 of LEA,
135-205 of length to origin of dorsal thong, 99-164 of anal
fin base, 296-460 of caudal peduncle); dorsal profile of head
angular, with a shallow concavity at the nape at all sizes (Fig.
19); distance from snout to rictus 61 1-1083 of snout length;
chin bulbous; snout sharply pointed; snout length 267-323
of head length, 935-1241 of interorbital distance; both jaws
edentulous in adults.

Eye diameter 90-136 of head length, 293-484 of snout
length, 314-500 of interorbital distance; interorbital distance
242-3 1 5 of head length; branchial opening 155-337 of depth
at nape, 153-271 of head length; distance from tip of snout
to vent 47-143 of LEA, vent shiftinganteriad with increasing
size (Fig. 1 2).

Anal fin base 778-951 of LEA; length of pectoral fin 885-
1000 of head length.

59-64 vertebrae to base of last anal fin ray.

Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus 17

Figure 19. Adontosternarchus sachsi (Peters) 205.0 mm TL, MBUCV-V-10441, Orinoco River, Cano Guine, Venezuela. A, entire fish; B,
close up of head.

Background color in alcohol pale tan to brown; sides and
back densely and almost uniformly peppered with brownish-
black chromatophores; chromatophores of mid-sides often
arranged as oblique bands along scale margins; mid-doral
pale stripe absent; dorsal thong with a hyaline ground color
but superficially covered with chromatophores; lateral line
sensory canal evident as a thin pale line on sides; superficial
chromatophores of lower sides sometimes forming ventro-
posteriorly oblique lines and underlain with numerous cor-
responding dark lines formed by deep chromatophores and
spaces between anal fin basals; anal fin membrane hyaline
but usually a few chromatophores present over rays; caudal
fin dusky at base and centrally, its distal end hyaline; pectoral
fins hyaline; top and upper sides of head pigmented as the
anterior part of the body; lower sides and undersurface of
head paler, with scattered chromatophores; tip of snout, mar-

gin of upper lip, tip of chin and tube of anterior naris almost
always immaculate.

DISTRIBUTION. Adontosternarchus sachsi is found in
the middle and lower parts of the Orinoco and Amazon rivers
(Fig. 17). Data on its detailed distribution in the Orinoco
Delta region was presented above in connection with A. de-
venanzii (p. 14 and Fig. 15).

ACKNOWLEDGMENTS

This work has been greatly improved through discussions
with Hector Lopez, Antonio Machado, and Edie Marsh. We
thank these people and the scientific party and crew of R/V
EASTWARD for their dedicated and able assistance on the
Orinoco Delta expeditions of 1978 and 1979.

Leslie Knapp and Frank Ferrari of the Smithsonian Ocean-

18 Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus

ographic Sorting Center provided much needed logistical
support for handling the Orinoco Delta collections. We thank
the following people and their institutions for loans of spec-
imens: W. Saul (ANSP), C.L. Smith (AMNH), W. Eschmeyer
and P. Sonoda (CAS), D.J. Stewart (FMNH), C. Swift
(LACM), J. Fernandez (MAC-PAY), K. Hartel (MCZ), N.
Menezes and H. Britski (MZUSP), R. Vari and J. Gomon
(USNM), R.M. Bailey and R.R. Miller (UMMZ). F. Kirsch-
baum provided a photograph and information on the type
of A. sachsi. K. Clark collected A. clarkae in the Rio Negro
and donated that material to MBUCV. R. West, on behalf
of the trustees of the Carnegie Museum, granted permission
to reproduce Ellis’s figure of A. balaenops. S. Murray assisted
with measurement and data management.

Much support for this work came from the National Sci-
ence Foundation grants DEB 77-14439 and DEB 80-22343.
We acknowledge use of R/V EASTWARD operated by Duke
University; the Duke National Oceanographic Facility was
supported in 1978 and 1979 by NSF, CG-QQQ05. Logistic,
personnel, and material support were provided by the Ven-
ezuelan Ministerio de Agriculture y Cria and Corporacion
Venezolana de Guayana. We thank the government and peo-
ple of Venezuela for permission to make collections and to
work in their country.

LITERATURE CITED

Bennett, M.V.L. 1971. Electric organs. In Fish physiology’,
ed. W.S. Hoar and D.J. Randall, vol. 5, pp. 347-491.
New York, Academic Press.

Cope, E.D. 1878. Synopsis of the fishes of the Peruvian
Amazon, obtained by Professor Orton during expedi-
tions of 1873 and 1877. Proceedings of the American
Philosophical Society 17:673-701.

Eigenmann, C.H. 1910. Catalogue of the freshwater fishes
of tropical and south temperate America. In Reports of
the Princeton University Expedition to Patagonia, 1896-
1899 3(4):375— 51 1.

Eigenmann, C.H., and W.R. Allen. 1 942. Fishes of western
South America. Lexington: University of Kentucky, xv
+ 494 pp.

Eigenmann, C.H. , and R.S. Eigenmann. 1891. A catalogue
of the fresh-water fishes of South America. Proceedings
of the United States National Museum 14:1-81.
Eigenmann, C.H. , and D.P. Ward. 1905. The Gymnotidae.

Proceedings of the Washington Academy of Sciences 7:

159-88.

Ellis, M.M. 1912. Order Glanencheli Family X. Gymno-
tidae, pages 422-42. In Eigenmann, C.H. The freshwater

fishes of British Guiana, including a study of the eco-
logical groupings of species and the relation of the fauna
of the plateau to that of the lowlands. Memoirs of the
Carnegie Museum 5:xii + 578 pp.

. 1913. The gymnotid eels of tropical America.

Memoirs of the Carnegie Museum 6: 109-95.

Fowler, H.W. 1915. Note on some gymnotid fishes. Copeia
no. 15, p. 2.

. 1939. A collection of fishes obtained by Mr. Wil-
liam C. Morrow in the Ucayali River Basin, Peru. Pro-
ceedings of the Academy of Natural Sciences Philadel-
phia 91:219-89.

. 1943. Los peces del Peru (Characininae to Gym-
notidae). Bo/ettn Museo de llistona Natural " Javier
Prado,” Universidad San Marcos, Lima 7(24-25):96-
124.

. 1945. Los peces del Peru. Cata/ogo sistemdtico de

los peces que habitan en aquas peruanas. Lima: Museo
de Historia Natural “Javier Prado,” Universidad Na-
cional Mayor de San Marcos. 298 pp.

. 1951. Os peixes de agua doce do Brasil. Arquivos

de Zoologia do estado de Sao Paulo 6(3):405— 628.

Ihering, R. von. 1907. Os peixes de agua doce do Brasil.
Revista Museu Paulista 7:258-336.

Lopez-Rojas, H., J.G. Lundberg, and E. Marsh. 1984. De-
sign and operation of a small trawling apparatus for use
with dugout canoes. North American Journal of Fish-
eries Management 4(3):33 1—334.

Mago-Leccia, F. 1967. Notas preliminares sobre los peces
de los llanos de Venezuela. Bolet'in de la Sociedad Ve-
nezolana de Ciencias Naturales Caracas 27(1 1 2):23 7—
63.

. 1970. Lista de los peces de Venezuela, incluyendo

un estudio preliminar sobre la ictiogeograffa de! pais.
Caracas: Oficina Nacional de Pesca, Ministerio de Agri-
culture y Cria. 238 pp.

Peters, W. 1877. Uber die von Hrn. Dr. C. Sachs in Ven-
ezuela gesammelten Fische. Monatsberichte Akademie
Wissenschaften Berlin 1877, pp. 469-73.

Rohl, E. 1942. Fauna descritiva de Venezuela. Caracas. 432

pp.

Sachs, C. 1879. Aus den Llanos. Schilderung enter natur-
wissenschaftlichen Reise nach Venezuela. Leipzig. 369

pp.

Schultz, L.P. 1949. A further contribution to the ichthy-
ology of Venezuela. Proceedings of the United States
National Museum 99(3235): 1-2 1 1.

Accepted 18 July 1984.

Contributions in Science, Number 358

Mago-Leccia, Lundberg, and Baskin: Adontosternarchus 19

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a*

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-% '-! •

N lumber 359
II April. 1985

BIOMECHANICAL IMPLICATIONS OF THE
VARIATION IN SMILODON ECTOCUNEIFORMS
FROM RANCHO LA BREA

Christopher A. Shaw and Antonia E. Tejada-FIores

mm

im'.

: ; V:

wBm

Up Si ■

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Printed at Alter. Press, Inc., Lawrence, Kansas.

BIOMECHANICAL IMPLICATIONS OF THE
VARIATION IN SMILODON ECTOCUNEIFORMS
FROM RANCHO LA BREA

Christopher A. Shaw and Antonia E. Tejada-Flores1

ABSTRACT. Of 5 1 7 Smilodon ectocuneiforms examined from the
late Pleistocene Rancho La Brea asphalt deposits, the typically hook-
like plantar process is unfused in 23.8 percent. The plantar tuberosity
of felid ectocuneiforms develops from a separate center of ossifica-
tion than that of the main body of the ectocuneiform. Tendons of
the M. tibialis posterior and M. peroneus longus. extensors of the
hind limb, interact with the plantar process of the ectocuneiform
and affect the development of this bone. The frequency of unfused
plantar tuberosities is greater in geologically younger localities at
Rancho La Brea and is correlative with an overall increase in Smi-
lodon body size. This increase in body mass through time placed
greater stress (produced by proportionately more powerful extensor
muscles) on the developing plantar process of larger Smilodon in-
dividuals, preventing fusion to the body of the ectocuneiform. Lever-
arm ratios, adjusted for body mass, indicate that Smilodon had
mechanically more powerful hind limbs than living lions and was
able to attack prey by stalking and leaping.

INTRODUCTION

Osteologically, Smilodon jloridanus is one of the most com-
pletely known extinct species from the late Pleistocene of
Rancho La Brea in southern Californa. Between 1913 and
1915, field parties from the Los Angeles County Museum of
Natural History (LACM) excavated extensively in what is
now Hancock Park. From 1029 to over 2000 estimated in-
dividuals of Smilodon are represented in the collections
(Marcus, 1960; Miller, 1968) housed at the George C. Page
Museum.

In a comparative osteologic study of the felids from Ran-
cho La Brea, Merriam and Stock (1932) noted two morpho-
logic varieties of Smilodon ectocuneiforms. The more com-
mon variety (Fig. 1A) has a hook-like plantar tuberosity,
fused to the body along a broad area; this process is often
truncated, the hook being reduced or absent in the less com-
mon variety (Fig. 2A). Merriam and Stock (1932) examined
1 20 ectocuneiforms and found that 70.8 percent bore a well-
developed tuberosity. This percentage is 76.2 in our study
of 517 specimens. Six of 11 (54.5%) Smilodon ectocunei-
forms from asphalt deposits of Talara, Peru, have well-de-

Contributions in Science, Number 359, pp. 1-8
Natural History Museum of Los Angeles County, 1985

veloped, fused plantar tuberosities (C.S. Churcher, written
comm.).

The plantar (posterior) end of truncated ectocuneiforms
also exhibits two distinct morphologies. In our sample of 1 23
specimens, 37 (30.1%) bear a facet (Fig. 2A, right) for ap-
parent articulation with a small nodule of bone, representing
the missing plantar process. The remainder have a roughened
surface for tendinous or ligamentous attachment similar to
that of the fully developed process; furthermore, nine of these
specimens (7.3%) exhibit unexplained pathologic bone growth.
Four of the five truncated ectocuneiforms from Talara bear
a nodular facet (C.S. Churcher, written comm.).

Until recently, the nodule of bone assumed to be associated
with the truncated Smilodon ectocuneiforms by Merriam and
Stock (1932) had not been recognized from Rancho La Brea.
Twenty specimens have now been identified in the collec-
tions. Seven (LACMHC 10791, 10792, 10795, 10805, 10806,
LACMRLP R3693 1 , R5 1053) are hook-hke in shape, iden-
tical in morphology to the fused plantar process of complete
ectocuneiforms (Fig. 3A); four (LACMHC 10791, 10792,
1 0805, LACMRLP R3693 1 ) bear an articular facet and three
do not. The other 13 specimens (LACMHC 10790, 10793,
10794, 10796 through 10803. 10807) do not have a articular
facet and are oval and globular in shape. The dorsal (anterior)
surface bears an oval rugosity where articulation with or
fusion to the body of the ectocuneiform would be expected
(Fig. 3B).

Ectocuneiforms of extinct felids (other than Smilodon) from
Rancho La Brea and those of several extant species from the
comparative collections of the American Museum of Natural
History, LACM, and the George C. Page Museum (RLB)
were examined. Of 40 Felts atrox specimens one was trun-
cated; the plantar projection is small and rounded, with a
smooth surface texture and no articular facet (Fig. 2C). Small

1. George C. Page Museum, 5801 Wilshire Boulevard, Los An-
geles, California 90036.

ISSN 0459-8113

2 cm

h-. di

Figure 1. Normally developed left ectocuneiforms of Smilodon and four species of Felis: proximal (left), lateral (center), and plantar (right)
views. A. Smilodon (LACMHC Q4602). B. F. atrox (LACMHC 10788). C. F. tigris (RLB 8). D. F. leo (LACM M 1 1 44). E. F. concolor (L ACM
Ml 456). Line drawing by Mark Hallett.

samples of extant felid species ( F . leo (20), F. tigris (2), F.
concolor (6), F. cattus (4), and Lynx rufus (4)) were also
examined; only one F. t/gra ectocuneiform was found to have
an unfused plantar process. The prominently hooked plantar
tuberosity of this specimen occurs as a separate accessory
bone articulated to the body of the ectocuneiform (Figs. 2B

and 3C). The articular facets are large and oval. Furthermore,
in this individual, the plantar tuberosity of the ectocuneiform
is fused on the left side but not on the right.

It is likely that the plantar process is an ontogenetic center
of ossification separate from the main body of the ectocu-
neiform. Thus far no specific mention of the occurrence of

2 Contributions in Science, Number 359

Shaw and Tejada-Flores: Smilodon Ectocuneiforms

2 cm

Figure 2. Truncated right ectocuneiforms of Smilodon and two species of Felis: proximal (left), medial (center), and plantar (right) views.
A. Smilodon (LACMHC Q4445). B. F. tigris (RLB 8). C. F. atrox (LACMHC 10789). Line drawing by Mark Hallett.

accessory or supernumerary bones associated with the ec-
tocuneiform in mammals has been found in the literature.
However, other accessory or supernumerary tarsal elements
are apparently common in some mammals (Davis, 1964;
Grant and Basmajian, 1 965). Two centers of ossification could
account for the asymmetry seen in the F. tigris individual,
with fusion taking place on the left side but not on the right.

ANATOMY

Descriptions of Smilodon and Felis atrox ectocuneiforms are
provided by Merriam and Stock (1932). The body of the
ectocuneiform in F. atrox is very similar to that of all extant
felids, but that of Smilodon is very different. The proximal
surface of the plantar process, between the body and hooked
tuberosity, forms a very broad and flat neck in Smilodon.
This neck consistently equals or exceeds the width of the
proximal articular surface nearest the area of fusion to the
body (Fig. 1A). In all other felids examined, this neck is
medio-faterally constricted and proximo-distally rounded
(oval to circular in cross section). It is smooth and Hares
transversely into the hooked or rounded area of tendon at-
tachment. In the unfused specimen of F. tigris, the neck is
flared and rugose around the area of articulation. The hook-
like character of the plantar tuberosity in Smilodon, men-

tioned by Merriam and Stock (1932), is highly developed in
F. tigris, less developed in F. leo and F. conco/or, and poorly
developed in F. atrox (Fig. 1 ). A summary of ectocuneiform
measurements is provided in Table 1.

The variation observed in Smilodon ectocuneiforms may
be attributed to muscular forces exerted on this bone. Two
muscles interact with the ectocuneiform in modern felids,
the M. tibialis posterior and the M. peroneus longus. These
have been reconstructed for F. atrox and Smilodon (Fig. 4).
The M. tibialis posterior originates on the entire medial sur-
face of the head of the fibula and between the oblique ridges
of the posterior surfaces of the tibia. The muscle ends in a
slender flat tendon that passes through the dorsal groove on
the medial surface of the tibia, then turns onto the plantar
surface of the foot and passes through a groove on the ventral
surface of the navicular. It inserts on the posterior surface of
the plantar tuberosity of the ectocuneiform and the lateral
tuberosity of the navicular. This muscle is an extensor of the
foot (Reighard and Jennings, 1951; Crouch, 1969). The M.
peroneus longus, a flexor of the foot, originates from two
areas of the fibula, the lateral surface of the head and the
proximal half of the lateral surface. This muscle terminates
in a slender tendon which passes through the groove on the
lateral surface of the lateral malleolus, then passes through
peroneal grooves of the calcaneum, cuboid, and ectocunei-

Contributions in Science, Number 359

Shaw and Tejada-Flores: Smilodon Ectocuneiforms 3

2 cm

fa. —n i —i— j

Figure 3. Unfused plantar processes of Smilodon and Felis tigris:
dorsal (left) and medial (right) views. A. Smilodon (LACMHC 10791),
hooked process with articular facet. B. Smilodon (LACMHC 10790),
ovoid process without articular facet. C. F. tigris (RLB 8). Line
drawing by Mark Hallett.

form. Large branches insert on the proximo-posterior bases
of the first and fifth metatarsal and slender branches insert
similarly on the other metatarsals (Reighard and Jennings,
1951; Crouch, 1969). In addition, the plantar tuberosity of
the ectocuneiform serves as the attachment of the plantar
navicularicuneiform ligaments (Davis, 1958).

BIOMECHANICS

Comparison of fore and hind limb size and strength in ma-
chairodonts has been discussed by many authors. Schaub
(1925) and Bohlin (1940, 1947) assert that the heavily de-
veloped fore limbs were not balanced by equally developed
hind limbs, therefore, Smilodon could not initiate a leaping
attack on a prey animal. However, Merriam and Stock (1932)
and Simpson (1941) state that the hind limb is weak only
relative to the fore limb and that this genus probably began
its attack by lunging or leaping.

A more exact comparison can be made by analyzing rel-
ative lever-arm ratios in the hind limbs of Smilodon and
true cats. Any mechanical action can be described by the
following equation,

F0 = F, x L/L0

where F0 = out-force, F, = in-force, L, = length of the in-
lever, and L0 = length of the out-lever (Hildebrand, 1974).

The out-force, in-lever, and out-lever may be calculated

or directly measured for extinct species. The out-force is
defined as the resistance (in this case, the body mass of the
animal) that must be overcome by a muscular system to
obtain momentum. Assuming only the hind limbs are em-
ployed to propel the animal forward and each foot bears an
equal load, F„ equals one half of the body mass (M/2). Body

Table 1. Summary of measurements (in mm) of normally developed
(normal) and truncated ectocuneiforms of Smilodon and Felis atrox
from Rancho La Brea. Abbreviations: N = sample size, OR = ob-
served range, x = mean, s = standard deviation, V = coefficient of
variation, Dr-PI = dorso-plantar, Px = proximal, Dt = distal. Mea-
surements taken after Merriam and Stock (1932).

N OR x s V

Smilodon

Normal

Dr-PI depth

182

31.4-45.6

39.0

2.539

6.510

Px-Dt length

187

15.1-23.6

18.8

1.491

7.931

Dt width

184

21.0-28.6

24.7

1.461

5.915

Truncated

Dr-PI depth

123

26.8-36.6

31.5

2.0 1 3

6.390

Px-Dt length

122

15.1-23.0

17.8

1.461

8.006

Dt width

122

20.8-27.9

25.1

1.384

5.514

Felis atrox

Normal

Dr-PI depth

38

43.6-58.0

51.1

3.589

7.023

Px-Dt length

39

20.6-33.5

26.9

2.297

8.539

Dt width

40

24.1-33.4

29.1

2.236

7.684

Truncated

Dr-PI depth

1

33.5

Px-Dt length

1

23.4

Dt width

1

26.2

Table 2. Measurements (in mm) of the minimum antero-posterior
diameter of Smilodon and Felis atrox femora. The pits are ranked
by order of descending mean weight (in kg) which is calculated using
the allometric constants of Alexander et al. (1979). Abbreviations
as in Table 1.

N

OR

X

s

x weight

Smilodon

Pit

3

121

27.4-36.6

31.9

1.921

154.3

61/67

92

25.4-36.6

31.5

1.851

149.0

4

78

27.7-35.1

30.8

1.636

139.9

77

61

26.0-33.6

30.5

2.003

136.2

13

36

26.2-32.9

30.4

1.461

134.9

Total sample

415

25.4-36.9

31.3

1.947

146.3

Felis atrox

Total sample

23

30.8-43.7

35.1

3.450

201.2

4 Contributions in Science, Number 359

Shaw and Tejada-Flores: Smilodon Ectocuneiforms

5 cm

Figure 4. Posterior view of distal hind limb showing the origin
and insertion of the M. tibialis posterior and M. peroneus longus in
Smilodon (left) and Felis atrox (right). Illustration by Mark Hallett.

mass was calculated from the minimum antero-posterior fe-
mur midshaft diameter of Smilodon and F. atrox (Table 2)
using the allometric constants of Alexander et al. (1979).
Lengths of the in-lever (distal end of calcaneum to calcaneal
pivot at tibia, L,) and out-lever (calcaneal pivot to end of
metatarsal III, LJ were calculated from mean lengths of the
calcaneum, navicular, ectocuneiform, and metatarsal III for
Smilodon, Felis atrox, and F. leo (Tables 3 and 4).

The in-force is produced by contraction of extensors of the
lower hind limb and will approximate the minimum amount
of contractile force required to overcome the inertia of the
body mass. These muscles include the M. gastrocnemius, M.
soleus, M. plantaris, M. peroneus brevis, and M. tibialis pos-
terior. The large M. gastrocnemius arises from the patella
and femoral sesamoids and inserts on the distal calcaneum.
The M. plantaris has a similar origin but crosses the tuber
calcis to insert on the plantar aponeurosis of the foot. The
other muscles originate along portions of the tibia and/or
fibula and insert on the distal calcaneum (M. soleus) or on
palmar areas of the foot. Using the mechanical equation
above, F, was calculated for Smilodon, Felis atrox, and F.
leo (Table 4).

The out-force will become greater as the in-force or in-
lever length is increased or as the out-lever length is de-
creased. The mean lever ratio (L,/L0) is nearly equal in Felis

atrox and F. leo. Because the L„ of Smilodon has been ap-
preciably shortened and the L, lengthened, this ratio is pro-
portionately much greater, increasing the potential out-force
capability of hind foot extension. Adjusting for allometric
differences in mass (M) between comparable species by using
M/2 = Fu and solving the mechanical equation gives the min-
imum amounts of in-force (F,) necessary to overcome inertia.
The smaller the value F,, the more powerful the muscular
contraction; the larger the value of F„ the more rapid the

Table 3. Proximo-distal length measurements (in mm) of hind foot
elements of Smilodon, Felix atrox, and F. leo. Statistics on metatar-
sal III of Smilodon taken from Menard (1947). Abbreviations: Pv-
Dt = pivot to distal, * = approximately. Other abbreviations as in
Table 1.

N

OR

X

s

V

Smilodon

Calcaneum

Total length

50

84.6-101.5

93.5

3.628

3.879

Pv-Dt length

50

20.8-28.8

24.4

1.999

8.188

Navicular

30

16.6-21.5

18.6

1.198

6.429

Ectocuneiform

187

15.1-23.6

18.8

1.491

7.931

Metatarsal III

759

*85-112

97.8

*4.7

*4.8

Felis atrox

Calcaneum

Total length

56

108.0-140.4

125.3

7.792

6.219

Pv-Dt length

56

34.1-48.9

40.9

3.579

8.744

Navicular

54

15.9-20.7

18.5

1.321

7.140

Ectocuneiform

39

20.6-33.5

26.9

2.297

8.539

Metatarsal III

49

132.0-157.8

145.7

7.292

5.004

Felis leo

Calcaneum

Total length

20

90.9-1 14.9

103.5

6.736

6.508

Pv-Dt length

20

28.9-37.7

34.0

2.708

7.965

Navicular

20

14.1-18.2

15.8

1.151

7.287

Ectocuneiform

19

18.7-23.0

21.1

1.259

5.969

Metatarsal III

19

1 12.4-139.0

126.8

8.107

6.393

Table 4. Lever-arm statistics used to determine relative hind limb
lever strength between Smilodon, Felis leo, and F. atrox where F„ =
F, x L/L„. Abbreviations: x = mean, L, = in-lever, L„ = out-lever,
M = mass, F„ = out-force, Fi = in-force. Weight of F. leo from Schal-
ler (1972).

Smilodon

Felis leo

Felis atrox

x L, (mm)

69.1

69.5

84.4

x L„ (mm)

159.6

197.7

232.9

ratio x L,/x L„

0.43

0.35

0.36

x M/2 (kg) (=FU)

72.9

81.0

100.6

minimum F,

169.5

231.4

279.4

Contributions in Science, Number 359

Shaw and Tejada-Flores: Smilodon Ectocuneiforms

Table 5. Ectocuneiform samples and radiocarbon dates on bone collagen from Pits 3, 4, 13, 61/67, and 77. The sites from Rancho La Brea
are ranked from oldest to youngest. Specimen data from Talara, Peru, provided by C.S. Churcher (written comm.) and radiocarbon dates from
Churcher (1966). Abbreviations: Lt = left, Rt = right, % = percent of sample from the site. Other abbreviations as in Table 1.

Pit

Normal

Truncated

N

OR dates

Total N

N

%

Lt

Rt

N

%

Lt

Rt

77

169

157

92.9

81

76

12

7.1

6

6

3

28,200-33,100

4

44

32

72.7

16

16

12

27.3

6

6

12

13,500-36,000

3

141

80

56.7

40

40

61

43.3

31

30

9

12,650-21,400

13

33

30

90.9

16

14

3

9.1

2

1

4

14,310-15,360

61/67

56

36

64.3

17

19

20

35.7

14

6

5

1 1,130-13,600

Other pits

74

64

79.7

37

22

15

20.3

7

8

-

-

Total sample

517

394

76.2

207

187

123

23.8

66

57

33

11,130-36,000

Talara

11

6

54.5

3

3

5

45.5

2

3

4

13,616-14,418

muscular contraction. Of the three species examined (Table
4), Smilodon exhibits the smallest F,, indicating a more pow-
erful hind limb than F. atrox and F. leo. Obviously, Felis
leo is adept at running, leaping and in-place wrestling when
pursuing prey. Smilodon, with an even more powerful foot
leverage system and less body mass, was able to propel itself
more powerfully than living lions. The robust forelimbs of
Smilodon appear to be yet more powerfully organized and
constructed than the hind. Therefore, although the hind limb
is “weaker” than the fore limb as Schaub (1925) and Bohlin
( 1 940, 1 947) state, and proportioned differently than in most
true cats (as argued by Merriam and Stock, 1932, and by
Simpson, 1941), both are clearly more powerful appendages
than those of large living cats. In addition, the minimum F,
(Table 4) shows that the La Brea lion (F. atrox) had a dis-
proportionately weaker (read “speedier”) foot leverage sys-
tem for its body mass than the smaller modern lion (F. leo).
Thus in the La Brea ecosystem, the roles of large felids were
played by the smaller, more powerfully legged sabertooth
and the larger, speedier-legged lion.

CHANGES THROUGH TIME

Morphologic change during Pleistocene time is documented
in many mammalian lineages. In some taxa (e.g., Felis onca),
size is the main difference between fossil and living forms
(Kurten, 1973). Many felid taxa exhibit a gradual overall size
reduction throughout the Pleistocene; in contrast, the size of
Smilodon increased (Kurten, 1965).

Samples of Smilodon ectocuneiforms are available from

Table 6. Chi-square values comparing frequencies of normally de-
veloped and truncated Smilodon ectocuneiforms from five pits at
Rancho La Brea. * = P < 0.05, ** = P < 0.01. Probabilities inter-
polated from Zar (1974).

Pit

3

61/67

4

13

61/67

0.36

4

**

*

13

**

**

**

77

**

♦ *

**

0.45

several radiocarbon dated localities (pits) at Rancho La Brea.
Most sites accumulated fossil specimens within one or two
brief, definable periods spanning about 3000 years or less.
These are fairly evenly spaced between 1 1,000 and 36,000
years B.P. (Akersten et al., 1983; Marcus and Berger, 1984).
The most reliable radiocarbon dates from Rancho La Brea
are those measured on bone collagen (Ho et al., 1969; Ak-
ersten et al., 1983). In many instances collagen dates can be
reliably correlated with stratigraphic depth within a single
pit. Specimens from Pits 3, 4, 13, 61/67, and 77 have been
radiocarbon dated and these sites contained the largest sam-
ples of ectocuneiforms (Table 5).

The frequency of normally developed (versus truncated)
ectocuneiforms is distinct in each of these five sites (Table
5). The chi-square values show significant differences be-
tween all sites except between Pits 3 and 61/67 and between
1 3 and 77 (Table 6). In addition, pits in which high frequency
of truncation is observed are roughly correlative to younger
radiocarbon horizons (Table 5 and Fig. 5) and increased body
mass (Table 2). As observed. Pits 3 and 61/67, with high

501

45-

<■_ Talara

40-

_ 61/67

35-

~o

30-

c

25-

£

20-

15-

10-

— 13

5-

i i i i 1 1 i — 'i 1 1 i

5 10 15 20 25 30 35 40

103 years BP

Figure 5. Change in frequency of Smilodon ectocuneiform trun-
cation through time of the samples from five localities at Rancho
La Brea and one from Talara, Peru. Dotted line represents the mean
ectocuneiform truncation for the total sample from Rancho La Brea.

6 Contributions in Science, Number 359

Shaw and Tejada-Flores: Smilodon Ectocuneiforms

truncation frequencies, are of relatively young radiocarbon
age and contain Smilodon with the largest mean body size.
The small sample of comparable age from Talara (Table 5)
also has a high frequency of ectocuneiform truncation. The
sample from Pit 4 was accumulated throughout the time span
represented at Rancho La Brea and, as expected, the ecto-
cuneiform truncation frequency and mean body mass are
most similar to the entire sample (Tables 5 and 6). Pit 77,
which contains animals with relatively small body mass and
low ectocuneiform truncation frequency, is one of the oldest
sites of fossil accumulation; Smilodon body mass and trun-
cation frequency is similar in Pit 13, although radiocarbon
dates indicate it to be a much younger deposit, comparable
in age to Pits 3 and 61/67.

The similarity between Pits 13 and 77 is unexpected. Ra-
diocarbon dates of specimens from these pits do not appear
inaccurate (Marcus, pers. comm.). The sample of Smilodon
from Pit 13 is the least variable in observed range and stan-
dard deviation of all samples from Rancho La Brea (Tables
2 and 5; Menard, 1947). Its temporal range is also the most
restricted, only spanning approximately 1000 years of ac-
cumulation (Table 5). Furthermore, the body mass calcula-
tions (Table 2) indicate a depositional episode biased toward
the entrapment of small individuals. When excavating Pit
13, Wyman (1914) remarked that the Smilodon materials
“appear to average small as compared to those from other
pits,” which is consistent with the low frequency of ectocu-
neiform truncation and the small body mass measured from
femoral diameters. Thus, the low frequency of ectocuneiform
truncation in Pit 1 3 near the terminal Pleistocene is due to
an accumulation bias selectively entrapping small individ-
uals of Smilodon.

DISCUSSION

Several factors may have contributed to the observed in-
crease in frequency of plantar process truncation in Smilodon
ectocuneiforms from Rancho La Brea. An important con-
sideration is the effect of allometric increase of muscle and
tendon size through time and the increased stress in the rel-
atively small area of insertion on (and interaction around)
the plantar tuberosity. As the M. tibialis posterior and M.
peroneus longus became more massive, so did their tendons.
The area of insertion of the M. tibialis posterior on normally
developed ectocuneiforms is about equal in Smilodon and
Felis atrox(a considerably larger animal); similarly, the groove
for the M. peroneus longus is the same absolute size in both
taxa (Figs. 1A and IE). These features imply that both mus-
cles in each taxon were of comparable size. However, the
proximo-distal length of the ectocuneiform is much shorter
in Smilodon (Table 1) and the neck connecting the plantar
process is flattened and less robust. If the plantar process and
body of the ectocuneiform were ontogenetically two centers
of ossification, movement of a large M. peroneus longus ten-
don and the contraction of a powerful M. tibialis posterior
would tend to continually pull the plantar process away from
the body, preventing fusion. When two centers of ossification
develop close together, an articular facet forms in the area

of syndesmotic attachment between the bone surfaces (Fig.
3A); if they ossify further apart the ovoid plantar process
(Fig. 3B) would likely function as a sesamoid within the M.
tibialis posterior tendon. With increase of Smilodon body
mass through time, the increased muscular stress on the plan-
tar process of the ectocuneiform could explain the observed
lower frequency of fusion between these centers of ossifica-
tion.

It is interesting to note that the Felidae possess the most
robust plantar tuberosity of all living mammals. Within this
family, its degree of development is roughly negatively as-
sociated with cursorial behavior. It is characteristic that less
cursorial felids have a large M. tibialis posterior (Ginsberg,
1961) and a robust plantar process with a pronounced hook
for its insertion (e.g., Felis tigris, F. onca)\ the most cursorial
felid ( Acinonyx ) has a reduced M. tibialis posterior and a
reduced plantar process, very similar to canids. F. leo and
F. atrox fall between these extremes. Morphologically, the
plantar process of Smilodon is most like that of F. tigris and
F. onca.

Of the large living cats, Felis onca is most similar to Smi-
lodon in limb proportions (Gonyea, 1976a). Though digiti-
grade, both taxa share several features of the hind limb (short,
stocky tibia and fibula, large M. tibialis posterior, and short,
slightly splayed metatarsals) characteristic of plantigrade car-
nivores (Ginsberg, 1961). Ginsberg (1961) noted that F. onca
rarely employs a “rapid pursuit phase” in prey capture, but
usually leaps with a single bound, overpowering its victim;
he suggests identical predatory behavior in machairodonts.
Gonyea ( 1 976b) stated that Smilodon probably used ambush
and stalking techniques to capture prey, immobilizing it with
the powerful front limbs. Similar limb proportions between
Felis onca and Smilodon have been interpreted as indicative
of habitat (Gonyea, 1976b); this similarity, however, is more
likely to reflect a common prey capture technique. This is
supported by the Smilodon limb mechanics presented here.

ACKNOWLEDGMENTS

We thank C.S. Churcher for providing data on those speci-
mens from Talara, Peru, that are housed at the Royal Ontario
Museum. George L. Callison contributed much time and
advice on the biomechanical aspects of this study. Compar-
ative material of Felis leo was made available by Daniel H.
Russell of the American Museum of Natural History. Wil-
liam A. Akersten, David E. Fortsch, John M. Harris, George
T. Jefferson, Leslie F. Marcus, and James P. Quinn read the
manuscript critically. Partial support of this research was
provided by the Los Angeles County Museum of Natural
History Foundation.

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dating of petroleum-impregnated bone from tar pits at
Rancho La Brea, California. Science 164:1051-52.

Kurten, B. 1965. The Pleistocene Felidae of Florida. Bul-
letin of the Florida State Museum 9:21 5-73.

. 1973. Pleistocene jaguars in North America. Com-

mentationes Biologicae Societas Scientiarum Fennica 62:
1-23.

Marcus, L.F. 1960. A census of the abundant large Pleis-
tocene mammals from Rancho La Brea. Contributions
in Science, no. 38, 1 1 pp. Natural History Museum of
Los Angeles County.

Marcus, L.F., and R. Berger. 1984. The significance of ra-
diocarbon dates for Rancho La Brea. Pages 159-83 in
Quaternary extinctions, ed. P. Martin and R. Klein. Tuc-
son: University of Arizona Press.

Menard, H.W., Jr. 1947. Measurements in length of the
metapodials of Smilodon. Original Data, Archives,
George C. Page Museum, Los Angeles, California.

Merriam, J.C., and C. Stock. 1 932. The Felidae of Rancho
La Brea. Carnegie Institution of Washington Publication
no. 422, 231 pp.

Miller, G..I. 1 968. On the age distribution o/’Smilodon cal-
ifornicus Bovard from Rancho La Brea. Contributions
in Science, no. 131, 17 pp. Natural History Museum of
Los Angeles County.

Reighard, J., and H.S. Jennings. 1951. Anatomy of the cat.
New York: Henry Holt and Company, 498 pp.

Schaller, G.B. 1972. The Serengeti lion. Chicago: Univer-
sity of Chicago Press, 480 pp.

Schaub, S. 1925. Ueber die Osteologie von Machaerodus
cultridens Cuvier. Eclogae Geologicae He/vetiae 19(1):
255-66.

Simpson, G.G. 1941. The function of saber-like canines in
carnivorous mammals. American Museum Novitates
1 130:1-12.

Wyman, L.E. 1914. Field notes. Rancho La Brea. Archives,
George C. Page Museum, Los Angeles, California.

Zar, J.H. 1974. Biostatistical analysis. New Jersey: Pren-
tice-Hall, Inc., 620 pp.

Accepted 26 October 1984.

8 Contributions in Science, Number 359

Shaw and Tejada-Flores: Smilodon Ectocuneiforms

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Number 360
11 April 1985

CONTRIBUTIONS IN SCIENCE

0—

'M

A REVIEW OF THE GORIID FISH GENUS MONISHIA SMITH, 1949,
FROM THE WESTERN INDIAN OCEAN AND RED SEA,

WITH DESCRIPTION OF A NEW SPECIES

Menachem Goren

ji|i|

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A REVIEW OF THE GOBIID FISH GENUS MONISH I A SMITH, 1949
FROM THE WESTERN INDIAN OCEAN AND RED SEA,
WITH DESCRIPTION OF A NEW SPECIES

Menachem Goren1

ABSTRACT. Five species have been included in the gobiid fish
genus Monishia Smith, 1949: M. william (Smith, 1947); M. oculata
Smith, 1959; M. sordida Smith, 1959; M. ochetica (Norman, 1927);
M. bulejiensis Hoda, 1983. These species are compared and re-
described. M. adamsoni. a new species from Pakistan, is described;
it is characterized by having 12 segmented dorsal rays, 22 pectoral
rays (upper 6 free) and 38-39 scales along the body.

INTRODUCTION

The genus Monishia Smith, 1 959, is known from the western
Indian Ocean and the Red Sea (Smith, 1959; Miller, 1973;
Goren, 1979); Smith (1959) included three Indian Ocean
species: M. william (Smith, 1947), type species; M. oculata
Smith, 1959; and M. sordida Smith, 1959. Other species
subsequently assigned to Monishia are: Gobius ocheticus
Norman, 1927, from the Suez Canal (Miller, 1973, 1978),
and Cabillus anchialinae Klausewitz, 1975, from the north-
ern Red Sea (Goren, 1979). The latter is synonymized with
M. ochetica in this paper. An additional species, M. bule-
jiensis Hoda, 1983, was described from Pakistan. Yet another
species discovered during a study of gobies from the Pakistan
coast, is herein described.

Smith ( 1 960) placed in Monishia the South African species
Gobius saldanha Barnard, 1927; Talbot and Penrith (1965)
transferred it to Ctenogobius. Winterbottom (1976), who
placed it in “the overladen catchall genus Gobius noted
that G. saldanha shows certain similarities to Monishia and
may ultimately be placed in that genus. Miller (1978) sug-
gested the removal of saldanha to Caffrogobius or Gobius;
Hoese and Winterbottom ( 1979) assigned the species to Ne-
matogobius. Further research is needed to clarify the taxo-
nomic affinities of saldanha; however, I do not consider the
species to be a member of Monishia. Miller (1978) suggested
that Ctenogobius godavariensts Rao, may belong to Moni-
shia; unfortunately, I could not obtain specimens to confirm
or reject this suggestion. Hoese and Winterbottom (1979)
provisionally included Acentrogobius simulus Smith, 1960,

Contributions in Science, Number 360, pp. 1-9
Natural History Museum of Los Angeles County, 1985

in Monishia; however, 1 do not place it there as it has a scaled
predorsal and the upper rays in its pectoral fins are not free.

The species of Monishia resemble each other in mor-
phology and color pattern, probably due to the similarity of
the habitats they populate. The fishes that belong to this genus
are small, usually 30-40 mm in total length, never longer
than 60 mm. The genus is characterized by having the upper
rays of the pectoral fins free (at least partly) and filamentous.
The pelvic fins are fully united with a well-developed fraen-
um. The caudal fin is rounded and shorter than the head.
Scales along the body 26-40. Most of the scales are ctenoid;
only the anterior ones are cycloid. The scales reach a line
from the upper base of the pectoral to a point on the first
dorsal fin base, usually its insertion. The head, predorsal, and
prepelvic regions are naked. Vertebrae (including ural cen-
trum) 27 (in two individuals of O. ochetica, 28). The lateral
line system consists of cephalic canals with pores and rows
of sensory papillae. The cephalic pore system consists of the
following: a pair of nasal pores; unpaired anterior and pos-
terior interorbital pores; pairs of supra-otic, anterio-otic, pos-
terio-otic, and intertemporal pores. All of these pores open
along a continuous canal. The anterior and posterior tem-
poral pores open in a separate canal, except in M. sordida in
which all of the above-mentioned pores are connected. Three
preopercular pores, on the posterior margin of the preopercle,
are connected by a canal. The sensory papillae are arranged
mostly in vertical rows with two horizontal rows on the cheek,
two on the opercle, and a discontinuous row above the oper-
cle and the pectoral base. The structure of the lateral line
system of each species is illustrated in the figures.

The genus Monishia is closely related to Hetere/eotris
Bleeker, 1874, and Coryoga/ops Smith, 1958. Each of the
three genera has a depressed head, unsealed head and nape,
4-6 vertical, and 1-2 horizontal rows of papillae on the cheek

1. Department of Zoology, George S. Wise Faculty of Life Sci-
ences, Tel Aviv University, Tel Aviv 69978, Israel.

ISSN 0459-8113

Table 1. A summary of selected diagnostic characters in six species of Monishia.

ochetica
n = 47

oculata
n = 6

sordida
n = 8

william
n = 10

ad am so ni
n = 3

bulejiensis
n = 5

Scale alone body

26-31

30-33

34-36

34-36

38-39

31-33

Transverse scale rows

8-9

1 1

12-13

9-10

10-1 1

9-10

Segmented rays in second
dorsal fin

10-1 1 (12)

9

1 1

(10) 11

12

1 1

Segmented rays in anal fin

9-10

8

9

9

9

9

Pectoral rays

16-18

17-18

22

22-23

22

20

Free pectoral rays

2

3-4

5

6-7

6

5

Gill rakers

1 + 1+5

1 + 1+4

1 + 1+6

1 + 1+4

1 + 1+6

1 + 1+6

Vertebrae (including
urostyle)

27 (28)

27

27

27

27

27

Scales on pectoral base

(n = 14)

-

+

-

-

-

arranged in a similar pattern. In addition the first gill slit of
Hetereleotris is closed by a membrane as in Monishia (at
least partly). The cephalic pore system of Corvoga/ops is
similar to that of Monishia william. Monishia differs from
Hetereleotris in possessing ctenoid scales (vs. cycloid or na-
ked body), in having gill rakers on the anterior gill arch (vs.
none) and in the interorbital pore being always unpaired holes
(vs. tubular, mostly paired pores). Corvogalops differs from
Monishia in having the first gill arch completely free, pos-
terior nostril a hole (vs. tubular) and pelvic fins completely
separated, with the tips of the rays free.

Little is known of the biology of the species of Monishia.
My specimens were all taken in shallow water, usually less
than 2 m deep, although a few specimens of M. ochetica were
collected at a depth of 8 m (in artificial reefs). The fish seems
to be resistant to extreme ecological conditions. M. ochetica
was found in Lake Timsah, Suez Canal, where temperatures
fluctuate from 14 to 26.5°C and salinity from 30 to 50 parts
per thousand (Miller, 1978). M. anchialinae was found for
the first time in the hyperhaline water bodies of the Cracks
at Ras Muhammad, Sinai Peninsula (Klausewitz, 1975). M.
william was found in brackish water at the mouth of Nora
River, southeast Africa, as well as in other localities and
habitats ( Winterbottom, 1976). The species that is described
here as new was found in tidal pools. The diet of M. william
consists of amphipods, isopods, polychaetes, and occasional
decapods and mollusks (Winterbottom, 1976). Very little is
known about the reproductive behavior of the genus. The
only available information refers to M. sordida: “. . . appar-
ently male, indicates buccal incubation, for its mouth is full
of eggs, similar to those found in a female” (Smith, 1959:
207).

METHODS AND MATERIALS

Specimens from the collections of the following institutions
were studied: British Museum (Natural History), London
(BMNH); Hebrew University of Jerusalem (HUJ); Natural
History Museum of Los Angeles County (LACM); J.L.B.

Smith Institute of Ichthyology, Grahamstown (RUSI);
Senckenberg Museum of Frankfurt (SMF); Zoological Mu-
seum of Tel Aviv University (TAU).

Counts of vertebrae were made from X-ray photographs
taken for all the listed material in this work excluding M.
ochetica, of which only 14 specimens were photographed
(TAU 7606, 5 spec.; TAU 6206 + TAU 621 1, 5 spec.; TAU
5592, 1 spec.; HUJ 7571, 3 spec.).

X-ray photographs were also used to verify the counts of
the spines and rays in the dorsal and anal fins.

All measurements and counts given in the text and the
tables are based on the material examined and listed. Mea-
surements are given in mm. In cases in which the total length
could not be measured due to a damaged caudal fin, TL is
followed by ?.

The lists of synonyms include only publications in which
the taxonomy of a species is discussed or changed, or in which
additional information about the species is provided.

COUNTS. A, D, P, V — anal, dorsal, pectoral, and pelvic
fins, respectively; the posterior bifid ray of second dorsal and
anal ray counted as one.

MEASUREMENTS. BD — body depth at the insertion of
the first dorsal fin; GR — number of gill rakers on anterior
gill arch; HL— head length from snout tip to upper attach-
ment of opercular membrane; LS— longitudinal scale counts
from upper attachment of the opercular membrane to the
end of the hypural; SL— standard length; TL— total length;
TR — number of transverse scale rows, counted from the or-
igin of second dorsal fin.

KEY TO SPECIES OF MONISHIA

1. Pectoral fin base with scales M. sordida Smith

-. Pectoral fin base naked 2

2. Dorsal segmented rays 9; anal segmented rays 8

M. oculata Smith

-. Dorsal segmented rays 10-12; anal segmented rays 9-10
3

2 Contributions in Science, Number 360

Goren: Indian Ocean Monishia

3. LS scales 38-39; segmented dorsal rays 12

M. adamsoni n. sp.

LS scales fewer than 37; segmented dorsal rays 10-11

(rarely 12, but then LS fewer than 32) 4

4. LS scales 26-30; pectoral fin rays 16-18

M. ochetica (Norman)

-. LS scales 32-36; pectoral fin rays 20-22 5

5. A tentacle on posterior part of upper margin of eye .

M. bulejiensis Hada

-. No tentacle on posterior part of upper margin of eye . .

M. william (Smith)

Species are also distinguished in Table 1.

SPECIES ACCOUNTS

Monishia adamsoni new species

Figures la, 2, 4a

MATERIAL EXAMINED. Holotype. LACM 38320-23,
TL 41.4 mm, SL 35.0 mm; Pakistan, Sind, small cove 4.8
km west ofNuclear Power Plant, 1 3. II. 1979, in shallow water
(to 2 m), rocky bottom with sandy pockets. Paratypes. TAU
8800, TL 42. 1 mm, SL 35.5 mm, data as for holotype; LACM
38310-26, TL 36.3 mm, SL 30.7 mm; Pakistan, Sind, Beluji
Point, 27.1.1979, tidepool, shallow water (to 60 cm), con-
glomerate bed with rocky rubble.

ADDITIONAL MATERIAL. RUSI 74-91 (part), I ex.,
TL 20.5 mm, SL 17.1 mm. South Africa, Beauchamp, 1 mile
E on road near Tacotet Bay, 0.3 miles E of St. Marie Bridge
at jutting head of the island, 6.111. 197 1 .

ETYMOLOGY. This species is named in honor of Thom-
as A. Adamson (formerly of the Natural History Museum of
Los Angeles County) for his genuine interest in Indo-West
Pacific fishes and his valuable contributions to ichthyology.

DIAGNOSIS. A Monishia with 1 2 segmented rays in sec-
ond dorsal fin; pectoral rays 22, the upper 6 free; filamentous;
scales along body 38-39; gill rakers 1 + 1+6. In addition
adamsoni has a distinctive arrangement of the rows of sen-
sory papillae on head (Fig. 4a). Comparisons are presented
in Table 1 .

DESCRIPTION (based on holotype and 2 paratypes). Body
elongate and compressed. Head depressed, its upper profile
moderately convex. Mouth almost horizontal. Maxillary
reaches level of front of eye. Three to four rows of teeth on
each jaw, outer teeth on both jaws and inner teeth on lower
jaw enlarged. A pair of curved canines on the sides of the
outer teeth row on lower jaw. Tongue truncate with a slight
median emargination. Posterior nostril a short tube at front
of eye. Anterior nostril a long tube, with a tentacle on top.
overhanging labial groove. Eyes of moderate size, interorbital
space narrow. Gill opening restricted, reaching to below pec-
toral base. Gill rakers short 1 + 1+ 6.

Body proportions presented in Table 2.

Fins. D VI, I 12; A I 9; P 22 — upper 6 rays free; the first
fin is little shorter than the second, which is about two thirds
of body depth. Pectoral fins rounded, not reaching level of
insertion of the second dorsal fin. Pelvic fins fully united with
a well-developed fraenum. The pelvics do not reach the anus.

Anal fin inserts below third segmented dorsal ray. Its height
about two thirds of body depth. Caudal fin rounded.

Scales. LS 38-39; TR 10-1 1; body scales ctenoid, except
for the three-four anterior transverse rows. Predorsal and
prepelvic regions, pectoral base, and mid belly completely
naked. Vertebrae (including ural centrum): 27.

Cephalic lateral line system. Position and number of pores
and papillae as in Figure 4a.

Color (preserved). Body and head brownish, covered with
irregular dark speckles. A dark blotch on the upper base of
the pectorals and a vertical dark band on the base of the
caudal fin. First dorsal fin with two oblique large dark bands.
The upper tip of the fin white. Second dorsal fin brownish.

Monishia bulejiensis Hoda, 1983

Figures lb, 3, 4b

Monishia bulejiensis Hoda, 1 983a: 1 1 1-1 15; 1983b: 143-147.

Type locality: Karachi coast, Pakistan.

MATERIAL EXAMINED. LACM 38310-27 (2 ex.), TL
35.5-38.1 mm, SL 29.8-31.5 mm; TAU 8801 (2 ex.), TL
32.7-36.5 mm, SL 27.2-30.1, LACM 38309-22, TL 31.3

fable 2. Measurements and body proportions of the types of Mon-
ishia adamsoni n. sp.

Holotype

(LACM

38320-23)

Paratype

(TAU

8800)

Paratype

(LACM

38310-26)

Total length (mm)

41.4

42.1

36.3

Standard length (mm)

35.0

35.5

30.7

Body depth (mm)

7.2

7.6

6.1

Head length (mm)

10.6

1 1.2

8.5

Head width (mm)

8.6

8.7

6.7

Standard length (% of TL)

84.5

84.3

84.5

Body depth (% of SL)

20.5

21.4

19.7

Head length (% of SL)

30.3

31.5

27.7

Head width (% of SL)

24.6

24.5

21.8

Head depth (% of SL)

20.0

19.1

18.6

Distance between snout and

first pectoral fin (% of SL)

39.1

37.4

38.1

Distance between snout and

second pectoral fin (% of SL)

57.4

58.3

57.0

Distance between snout and

anal fin (% of SL)

62.8

61.9

62.3

Eye diameter (% of SL)

8.0

8.7

8.4

Longest pectoral ray (% of SL)

19.1

19.1

18.2

Longest spine in first dorsal

fin (% of SL)

11.1

12.9

12.0

Longest ray in second dorsal

fin (% of SL)

12.8

14.6

13.7

Longest anal ray (% of SL)

14.0

14.3

13.0

Head width (% of HL)

81.1

77.6

78.8

Eye diameter (% of HL)

26.4

27.6

26.8

Contributions in Science, Number 360

Goren: Indian Ocean Monishia 3

Figure 1. Photographs of six species of Monishia. a) M. adamsoni n. sp. Holotype LACM 38320, SL 35.0 mm. Sind; Pakistan, b) M.
bulejiensis LACM 38310, SL 31.5 mm. Sind; Pakistan, c) M. ochetica TAU 7607, SL 45.7 mm. Ras Muhammad, Sinai Peninsula, d) M.
oculata RUS1 830 (Paratype), SL 20.5 mm. Baixo Pinda. e) M. sordida RUSI 823 (Paratype), SL 37.5 mm. Inhaca Islands. 0 M. william
TAU 8803 (formerly RUSI 74-329), SL 32.5 mm. Coffee Bay. Photographs by L. Maman.

mm, SL 26.3 mm; LACM 38210 and TAU 8801 were col-
lected in Pakistan, Sind, Beluji Point, 27.1.1979, in shallow
water (to 60 cm) (tidepool), conglomerate bed with rocky
rubble. LACM 38309 was collected in Pakistan, Sind, Beluji
Point (300 m WNW of the point), shallow water (tidepool —
0.15 cm), conglomerate bed, 27.1.1979.

DIAGNOSIS. A Monishia with 1 1 segmented rays in sec-
ond dorsal hn and 9 in anal fin; pectoral rays 20, the upper
6 free and filamentous; scales along body 31-33; transverse
scale rows 9-10; a small tentacle on the upper posterior edge
of the eye. Possession of this tentacle distinguishes M. bu-
lejiensis from all known Monishia. In addition M. bulejiensis
has a distinctive arrangement of the rows of sensory papillae
(Fig. 4b). Comparisons are presented in Table 1.

DESCRIPTION (based on all specimens listed). Body
elongate and compressed. Head depressed, its upper profile
convex. Mouth oblique. Maxillae reach to level of mid eye.
On the upper posterior part of eye, just within the margin,
is a short cylindrical pointed tentacle, an outgrowth of the
sclerotic coat, its length equal to the length of the anterior
nostril. Each jaw with 3-4 rows of teeth, the outer teeth on
each jaw enlarged. No canines. Tongue truncate. Gill opening
restricted, reaching to below pectoral base. Gill rakers 1 + 1+ 6.
Vertebrae (including ural centrum): 27.

Cephalic lateral line system. Position and number of pa-
pillae as in Figure 4b.

Body proportions presented in Table 3.

Fins. D VI, I 1 1 ; A I 9; P 20, upper 5 rays free. First dorsal

4 Contributions in Science, Number 360

Goren: Indian Ocean Monishia

Figure 2. Monishia adamsoni n. sp. Holotype LACM 38320, SL 35.0 mm. Sind; Pakistan. Drawing by W. Ferguson.

tin little shorter than second dorsal fin, which is about two
thirds of body depth. Anal fin inserted below second dorsal
segmented ray. Pectoral fin rounded, reaching to below sixth
dorsal spine. Pelvic fins fully united with a well-developed
fraenum. Caudal fin rounded.

Scales. LS 31-33; TR 9-10. The body is covered with
ctenoid scales except those on belly and three-four anterior
rows on body which are cycloid. No scales on the median
line of the belly, on predorsal, pectoral base, and prepelvic
regions.

Color (preserved). Brownish, irregular dark speckles. A
series of light spots along the midlateral scale row. First dorsal
fin dark with upper margin white (about one quarter of its
height). Second dorsal fin and anal fin dark with a light line
along their edges. Pectoral fin dusky. Caudal fin with irregular
vertical dark rows.

Monishici ochetica (Norman, 1927)

Figures lc, 4c

Gobius ocheticus Norman, 1927:381, figs. 92-93. Type lo-
cality: Suez Canal.

Pomatoschistus (Ninnia) ocheticus: De Buen. 1930:132.

Coryphopterus ocheticus: Smith, 1959:21 1, pi. 93H.
Monishia ochetica: Miller, 1973:501; Miller, 1978:38-58.
CabiUus anchialinae Klausewitz, 1975:203-207.

Monishia anchialinae: Goren, 1979:46-48.

MATERIAL EXAMINED. BMNH 1925.9.19.114-123,

(syntypesof G. ocheticus), 7 ex., TL 18.2-32.4 mm, SL 1 6.2—
28.2 mm, Suez Canal, Lake Timsah, 1924; BMNH
1 925. 1 2.3 1.54-58 (syntypes of G. ocheticus ), 4 ex., TL 21.5-
33.0 mm, SL 1 8.2-27.0 mm, Suez Canal, 1924; SMF 13229
(holotype of Cabillus anchialinae), TL 41.1 mm, SL 33.4
mm; SMF 13230-13232 (paratypesof C. anchialinae), 3 ex.,
TL 26.7-34.5 mm, SL 23.2-28.6 mm, Sinai Peninsula, Cracks
at Ras Muhammad (detailed description of the biotope in
Por and Tsurnamal, 1973), VI. 1972; TAU 7606, 5 ex., TL
33.1-51.0 mm, SL 28.0-45.7 mm. Cracks at Ras Muham-
mad, 1 6.X. 1 969; 26 ex., around Sinai Peninsula (detailed list
in Goren, 1979, as Monishia anchialinae), TL 24.0-47.0
mm, SL 20.0-38.4 mm; HUJ E 62/532, Southern Red Sea,
Nocra, 1 ex., TL 24.0 mm, SL 20.0 mm, 23.III. 1 962.

DIAGNOSIS. A Monishia with 10-11 segmented rays in
second dorsal fin and 9-10 in anal fin; pectoral rays 16-18,
upper 2 partly free; scales along body 23-3 1 ; transverse scale

Table 3. Selected body proportions of M. ochetica, M. oculata, M. sordida, M. william, and M. hulejiensis.

ochetica
n = 47

oculata
n = 6

sordida
n = 8

william
n = 10

bulejiensis
n = 5

Total length (mm)

24.0-51.0

15.5-25.6

32.7-43.3

27.8-41.2

31.3-38.1

Standard length (mm)

20.0-45.7

12.8-21.3

27.9-37.5

22.9-34.0

26.3-31.5

Standard length (% of TL)

81.0-84.2

81.4-83.2

82.2-86.8

81.5-83.9

82.5-84.0

Head length (% of SL)

27.8-30.5

27.9-29.1

29.6-32.9

29.4-30.2

28.5-30.4

Body depth (% of SL)

18.1-23.2

15.9-16.7

19.7-20.6

18.3-21.1

17.6-20.8

Distance between snout and

first dorsal fin (% of SL)

34.2-36.8

35.4-37.4

35.3-36.5

36.1-37.6

35.4-39.0

Distance between snout and

second dorsal fin (% of SL)

52.3-56.0

54.3-56.3

54.6-56.6

54.4-58.2

52.2-56.2

Distance between snout and

anal fin (% of SL)

52.7-56.2

54.6-56.3

56.6-58.6

54.4-57.3

57.0-60.4

Head width (% of HL)

61.1-71.2

51.2-53.0

70.2-71.4

68.5-71.4

71.4-75.0

Eye diameter (% of HL)

25.4-30.1

25.0-26.0

25.0-26.1

24.2-25.0

25.0-27.5

Contributions in Science, Number 360

Goren: Indian Ocean Monishia 5

Figure 3. Monishia bulejiensis LACM 38310, SL 31.5 mm. Sind; Pakistan. Drawing by W. Ferguson.

rows 8-9; gill rakers 1 + 1+ 7. Comparisons are presented in
Table 1.

DESCRIPTION (based on the above listed material). Body
elongate and compressed. Head slightly depressed. Mouth
small, oblique. Three-four rows of teeth on each jaw; outer
teeth larger, inner a little curved backward. Tongue bilobed.
Posterior nostril a very short tube, in front of eye. Anterior
nostril tubular, close to upper lip. Interorbital space narrow,
about one half pupil diameter. Gill opening restricted. Gill
rakers 1 + 1 +5-7, relatively short.

Cephalic lateral line system. Position and number of pores
and papillae as in Figure 4c.

Body proportions presented in Table 3.

Fins. D VI, I 10-1 1; A I 9-10; P 16-18, 2 upper rays partly
free. In the isolated population at the Cracks at Ras Muham-
mad (the type locality of M. anchialinae) two specimens were
found with seven spines in first dorsal hn and 12 segmented
dorsal rays. First dorsal hn high relative to that of the other
Monishia species. Longest dorsal spines equal to body depth;
second dorsal tin reaches almost the same height. Anal hn
inserts below second dorsal segmented ray. Pectoral fins
rounded, each to below the insertion of the second dorsal
hn. Pelvic hns fully united. Fraenum weak. Caudal hn round-
ed.

Scales. LS 26-31; TR 8-9. Body covered with ctenoid
scales except those on belly which are cycloid. The scales
reach a line from the insertion of the first dorsal hn to the
upper part of pectoral base. No scales on predorsal and pre-
pelvic areas, and none on pectoral base.

Vertebrae. 27 including ural centrum (28 in 2 specimens
from the Cracks at Ras Muhammad, not those with 7 spines
or 12 segmented dorsal rays).

Color. Body and head brownish. Dark blotches along me-
dian line of the body. Irregular darker bands on body. Head
darker than body. Back of the body and the upper part of
head darker than the rest of the body. First dorsal hn with
two large diagonal black bands. In several specimens the hrst
dorsal hn is uniformly dark with its tip white. Second dorsal
hn and anal hn dusky with white edges. Pelvic hns dark.
Caudal hn with vertical irregular lines of dark spots.

REMARKS. In a previous paper (Goren, 1979) I noted
that M. anchialinae is in fact a synonym of M. ochetica, but
I left the two species separate as I had some hestitation that

arose from apparently contradictory information concerning
the habitats of the two nominal species. A further study in
which fresh material from the Cracks at Ras Muhammad
was included, as well as many observations in the Red Sea
during the last four years, settled the question. These hsh
were observed always on or near hard substratum which in
most of the occasions was located in sandy areas. In certain
cases it was a small coral knoll, but usually the hsh were
observed in or near small coral heads, mollusk shells, and
even artificial objects such as bottles and tins. The hsh searched
for food on the sand in close proximity to the hard substratum
and hid in holes when disturbed. I conclude that when spec-
imens of M. ochetica were collected from what seemed to be
soft bottom, hard substratum was actually part of the habitat.
Thus, on the basis of these new observations and additional
examinations of material, I regard M. anchialinae ns a junior
synonym of M. ochetica.

Monishia oculata Smith, 1959

Figures Id, 4d

Monishia oculata Smith, 1959:206. Type locality: Mahe;

Miller, 1978:56.

MATERIAL EXAMINED. RUSI 830, paratypes, 3 ex.,
TL ?, SL 15.3-20.5 mm, Baixo Pinda, 10. IX. 1956; RUSI
74-95 (part), 3 ex., TL 15.5-25.9 mm, SL 12.8-21.3 mm,
Beauchamp, 1 mile E on road near Jacotet Bay, 0.3 miles E
of St. Marie Bridge at jutting head of island, 6. III. 1971.

DIAGNOSIS. A Monishia with a wide dark band from
eye to lower edge of preopercle. Second dorsal hn with 9
segmented rays. Anal hn with 8 segmented rays; pectoral rays
17-18, upper 3-4 free; scales along body 30-33; transverse
scale rows 1 1; hrst gill slit closed by a membrane. M. oculata
can be distinguished from all its congeners by a diagonal dark
band below eye. Comparisons are presented in Table 1.

DESCRIPTION (based on the above listed material). Body
elongate and compressed. Head depressed. Mouth little
oblique. Maxillae to level of mid eye. Three to four rows of
teeth on each jaw. Outer teeth enlarged. The two lateral pairs
of teeth in the inner row of the lower jaw are caniniform.
Tongue bilobed. Posterior nostril a short tube, in front of
eye. Anterior nostril tube above lip. No flaps on nostrils. Gill

6 Contributions in Science, Number 360

Goren: Indian Ocean Monishia

Figure 4. Lateral line sensory papillae and canal pores in six species of Monishia. a) M. adamsoni n. sp. Holotype LACM 38320. Sind;
Pakistan, b) M. bulejiensis LACM 38310. Sind; Pakistan, c) M. ochetica TAU 7606, Ras Muhammad, Sinai Peninsula, d) M. oculata RUSI
830 (Paratype). Baixo Pinda. e) M. sordida RUSI 823 (Paratype). Inhaca Islands. 0 M. william TAU 8803 (formerly RUSI 74-329). ColTee
Bay. Drawing by W. Ferguson.

Contributions in Science, Number 360

Goren; Indian Ocean Monishia 7

opening restricted. First gill slit closed by a membrane. Gill
rakers 1 + 1+ 4, very short.

Vertebrae (including ural centrum). 27.

Cephalic lateral line system. As shown in Figure 4d.

Body proportions are presented in Table 3.

Fins. D VI, I 9; A I 8; P 17-18, upper 3-4 rays free. Dorsal
fin height less than body depth. Pectoral fins rounded, reach-
ing to below second dorsal segmented rays. Upper pelvic rays
free. Pelvic fins fully united, almost reaching the anus. Frae-
num developed. Caudal fin rounded.

Scales. LS 30-33; TR 1 1 . Ctenoid scales on body, reaching
to a line from second dorsal spine to upper part of pectoral
base. No scales on mid belly, pectoral base, prepelvic and
predorsal regions.

Color (preserved). Body yellowish with four vertical wide
and irregular brown bands; that on caudal base much darker.
A dark wide band runs from eye down and backward to
preopercle margin. Two oblique dark bands on first dorsal
fin. Dark spots form irregular vertical lines on second dorsal
and anal fins. Pelvic fins colorless to dark. Caudal fin col-
orless.

Monishia sordida Smith, 1959

Figures le, 4e

Monishia sordida Smith, 1959:206-207. Type locality; In-

haca; Miller, 1978:55-56.

MATERIAL EXAMINED. RUSI 823, paratypes, 3 ex.,
TL 38. 1-43.3 mm, SL 32.3-37.5 mm, Mozambique, Inhaca
Islands, VIII. 1948; TAU 8802 (out of RUSI 3018), 5 ex.,
TL 32.7-41.7 mm, SL 27.9-34.0 mm, Mozambique, Lou-
renco Marques, Ponte Maone, X.1953.

DIAGNOSIS. A Monishia with 2-3 vertical rows of scales
on pectoral base; scales along the body 34-36, transverse
scale rows 12-13; segmented rays in second dorsal fin 1 1 and
in anal fin 9; pectoral rays 21-22, the upper 5 free. Monishia
sordida differs from all its congeners in having scales on
pectoral base. Comparisons are presented in Table 1.

DESCRIPTION (based on the above listed material). Body
elongate and compressed. Head depressed. Snout convex,
little shorter than eye diameter. Mouth oblique. Maxillae
reach to level of mid eye, 3-4 rows of teeth on each jaw, the
outer enlarged. At the middle of the inner row of the upper
jaw two large canines directed backward. Two to three curved
canines on each side of inner row on lower jaw. Tongue
slightly emarginate. Posterior nostril a short tube at the front
of eye. Anterior nostril, a tube with a Hap, hanging above
lip. Gill opening restricted. Gill rakers relatively long, 1 + 1 +6.

Vertebrae (including ural centrum). 27.

Cephalic lateral line system. As in Figure 4e.

Body proportions presented in Table 3.

Fins. D VI, I 1 1; A I 9; P 22 (5 free). The two dorsal fins
are the same height, which is about two thirds of body depth.
Pectoral fins rounded, reaching to below insertion of second
dorsal fin. Pelvic fins fully united with a weak fraenum. The
pelvics reach 65-80 percent of the distance between the anus
and the pelvic fin base. Caudal fin rounded.

Scales. LS 34-36; TR 12-13. The body covered with scales

posteriorly from a line from the first dorsal fin insertion to
the upper base of the pectoral fin. Scales in the anterior 2-3
transverse rows are cycloid, the rest are ctenoid. Belly and
pectoral base covered with cycloid scales. No scales on pre-
pelvic and predorsal regions.

Color (preserved). Body brownish. A black blotch on the
upper part of the pectoral base. Two wide diagonal dark
bands on first dorsal fin. Dark dots form diagonal lines on
the second dorsal fin and vertical lines on the caudal fin.

REMARKS. Winterbottom (1976) found prepelvic scales
in some specimens of M. sordida “(Not in all individuals),”
but I found none.

Monishia william (Smith, 1947)

Figures If, 4f

Bathygobius william Smith, 1947:340. Type locality: Xora

River Mouth, Transkei (lectotype by Winterbottom, 1976).
Monishia william: Smith, 1959:206-207; Winterbottom,

1976:1-1 1; Miller, 1978:49-56.

MATERIAL EXAMINED. RUSI 74-348, 1 ex., TL 39.0
mm, SL 32.5 mm, Chaka's Rocks, Natal, 3.IX.1974; RUSI
77-12, 2 ex., TL 34.1-37.5 mm, SL 27.8-30.5 mm, Port
Edward, 3 1 .V. 1 977; RUSI 74-322, 2 ex., TL 27.8-28.8 mm,
SL 22.9-23.4 mm, Tshani, Transkei, VIII. 1974; TAU 8803
(formerly RUSI 74-329), 5 ex., TL 32.5-41.2, SL 27.0-34.0
mm. Coffee Bay, 6. VIII. 1974.

DIAGNOSIS. A Monishia with 1 1 (rarely 10) segmented
rays in second dorsal fin and 9 in anal fin. Scales along body
34-36; transverse scale rows 9-10; pectoral fin with 22-23
rays, upper 6-7 free; first gill slit closed by a membrane; gill
rakers very short (1 + 1+4). Comparisons are presented in
Table 1.

DESCRIPTION (based on the above listed material). Body
elongate and compressed. Head slightly depressed. Snout
oblique-convex. Posterior nostril a short tube in front of eye.
Anterior nostril a moderate tube (about one third diameter
of pupil) with a skin flap at its hind margin. Mouth little
oblique. Maxillae reach to below mid eye. Two to three rows
of teeth on both jaws. Outer teeth on upper and lower jaws
enlarged and pointed. A pair of canines at the side of the
lower jaw. Tongue rounded. Interorbital space narrow, about
one third diameter of pupil. Gill opening restricted, reaching
lower part of pectoral base. First gill slit closed by a mem-
brane. Gill rakers short, I + 1 +4.

Vertebrae (including ural centrum). 27.

Cephalic lateral line system. As in Figure 4f.

Body proportions presented in Table 3.

Fins. D VI, I 11 (10 in one); A I 9; P 22-23, the upper 6
or 7 rays free. Dorsal fins of equal height; longer spines and
rays reach about two thirds of body depth. Second dorsal fin
inserting above anus. Anal fin inserting below second dorsal
segmented ray, its height about two thirds of body depth.
Pectoral fins rounded, reaching to below insertion of second
dorsal fin. Pelvic fins fully united, fraenum well developed.
Caudal fin rounded.

Scales. LS 34-36; TR 9-10. Body covered with scales to
a line from insertion of first dorsal fin to upper base of pec-

8 Contributions in Science, Number 360

Goren: Indian Ocean Monishia

toral fin. Scales in anterior 3-4 transverse rows and those on
belly are cycloid, the rest are ctenoid. Mid belly, pectoral
bases, and predorsal and prepelvic regions unsealed.

Color (preserved). Body brownish with 4-6 irregular dark
vertical bands. Black blotches along median line of body.
Dark vertical half-moon blotch on the base of the caudal fin.
A dark blotch on the upper part of the pectoral base; first
dorsal fin with two diagonal wide dark bands. Rows of black
spots form vertical lines on the caudal fin and diagonal lines
on second dorsal and anal fin.

Remarks. The counts presented in this work are very close
to those given in the original description of M. william (Smith,
1947) but differ from those given in subsequent publications
(Smith, 1959, 1960). The reason for this discrepancy is the
inclusion of specimens of Bathygobius fuscus in the material
of M. william as found by Winterbottom (1976).

ACKNOWLEDGMENTS

1 am grateful to T.A. Adamson (formerly LACM) for pro-
viding the material from Pakistan and to Professor M.M.
Smith (RUSI) for providing material from South Africa. Fi-
nally, I thank Dr. A. Freidberg for reviewing this paper.

LITERATURE CITED

De Buen, F. 1930. Sur une collection de Gobiinae pro-
venant du Maroc. Essai de synopsis des especes de l'Eu-
rope. Bulletin de la Societe des Sciences Naturelles du
Maroc 10:120-47.

Goren, M. 1979. The Gobiinae of the Red Sea (Pisces:

Gobiidae). Senckenbergiana Biologica 60( 1/2): 1 3—64.
Hoda, S.M.S. 1983a. A new species of gobiid fish Monishia
bulejiensis (Family: Gobiidae) from the Coast of Paki-
stan. Biologia 29(1): I 14-15.

. 1983b. A new species of gobiid fish Monishia bu-
lejiensis (Teleostei: Gobiidae) from the Karachi coast.
Indian Journal of Fisheries 30( 1 ): 143-47.

Hoese, D. F., and R. Winterbottom. 1979. A new species

of Lioteres (Pisces, Gobiidae) from Kwazulu, with a re-
vised checklist of the South African gobies and com-
ments on the generic relationship and endemism of East-
ern Indian Ocean gobioids. Royal Ontario Museum Life
Sciences. Occasional Papers 31:1-13.

Klausewitz, W. 1975. Cabillus anchia/inae, eine neue
Meergrundel von der Sinai-Halbinsel (Pisces: Gobiidae:
Gobiinae). Senckenbergiana Biologica 56(4/6):203-07.

Miller, P.J. 1973. Gobiidae. In Check list of the fishes of
the North-Eastern Atlantic and of the Mediterranean 1 :
483-515.

. 1978. The systematic position and origin of Gobius

ocheticus Norman, 1927, from the Suez Canal. Zoolog-
ical Journal of the Linnean Society 62:39-58.

Norman, J.R. 1927. Zoological results of the Cambridge
Expedition to the Suez Canal, 1924. Fishes. Transac-
tions of the Zoological Society of London 22:375-89.

Por, F.D., and M. Tsurnamal. 1973. Ecology of the Ras
Muhammad Cracks in Sinai. Nature 24 1 (5384):43-44.

Smith, J.L.B. 1947. Brief revision and new records of South
African marine fishes. Annals and Magazine of Natural
History 14(1 1):335— 46.

. 1959. Gobioid fishes of the families Gobiidae, Peri-

ophthalmidae, Trypauchenidae, Taenioididae and
Kraemeriidae of the Western Indian Ocean. Ichthyolog-
ical Bulletin 13:185-225.

. 1960. Fishes of the family Gobiidae in South Af-
rica. Ichthyological Bulletin 18:299-314.

Talbot, F.H., and M.L. Penrith. 1956. Ctenogobius cloatus
Smith, 1960, a synonym of Ctenogobius saldanha (Bar-
nard, 1927). Annals of the South African Museum 48:
189-93.

Winterbottom, R. 1976. Notes of South African gobies
possessing free upper pectoral fin rays (Pisces: Gobiidae).
J.L.B. Smith Institute of Ichthyology. Special Publica-
tion 1 6: 1-1 1 .

Accepted 5 October 1984.

Contributions in Science, Number 360

Goren: Indian Ocean Monishia 9

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Number 361
13 June 1985

CONTRIBUTIONS IN SCIENCE

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THE SYSTEMATICS OF THE HYLAEINE BEES (HYMENOFI ERA: COLLETIDAE)
OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION: THE GENERA AND
SUBGENERA WITH REVISIONS OF THE SMALLER GROUPS

Roy R. Snelling

Natural History Museum of Los Angeles County • 900 Exposition Boulevard * Los Angeles, California 90007

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THE SYSTEMATICS OF THE HYLAEINE BEES (HYMENOPTERA: COLLETIDAE)
OF THE ETHIOPIAN ZOOGEOGRAPHICAL REGION: THE GENERA AND
SUBGENERA WITH REVISIONS OF THE SMALLER GROUPS

Roy R. Snelling1

ABSTRACT. The genera and subgenera of hylaeine bees of the
Ethiopian Region are characterized and separated by a key; pertinent
morphological features are illustrated. Within Hylaeus four subgen-
era are recognized: Deranchylaeus Bridwell and Metylaeus Bridwell,
as well as two new subgenera, Alfkenylaeus and Cornylaeus. Not-
hylaeus Bridwell (=Anylaeus Bridwell n. syn.) is regarded as a genus
apart from Hylaeus. Two new genera are described: Calloprosopis
in Kenya and Psilylaeus in Cape Province. All groups except De-
ranchylaeus and Nolhylaeus are revised in this part. Prosopis albo-
nasata Strand is a synonym of the Palaearctic species, H. signatus
(Panzer) and is probably incorrectly cited from “Kapland.” The
following species, described as hylaeines, are all allodapine antho-
phorids: Prosopis gracilis Bingham, P. pernix Bingham, P. quadri-
lineata Cameron, P. 5-lineata Cameron, and P. sandaracta Bing-
ham.

INTRODUCTION

This is the first of three parts treating the hylaeine bees of
the Ethiopian zoogeograph ical region. For purposes of this
study, the Ethiopian zoogeographical region encompasses all
of the African continent below the Sahara Desert, including
the Cape Region, but excluding the Malagasy Region. This
introductory part includes keys to the genera and subgenera,
revisions of all groups except Hylaeus, subgenus Deranchy-
laeus, and the genus Nolhylaeus. The second part will revise
the genus Nothylaeus and the final part will treat the species
here assigned to the subgenus Deranchylaeus of Hylaeus.

Early work on these bees consisted of descriptions of var-
ious species in papers by J.D. Alfken, P. Cameron, T.D.A.
Cockerell, H. Friese, F. Smith, and E. Strand (see Literature
Cited); these species were described, for the most part, under
the old generic name Prosopis, though some of the works by
Cockerell employed Hylaeus. In 1919 J.C. Bridwell attempt-
ed to organize the, by then numerous, species into genera
and subgenera. He recognized three genera: Nothylaeus, Me-
tylaeus, and Hylaeus. Nothylaeus was further divided into
two subgenera, Nothylaeus and Anylaeus. Those species as-
signed to Hylaeus were all placed in his new subgenus De-
ranchylaeus.

Contributions in Science, Number 361, pp. 1-33
Natural History Museum of Los Angeles County, 1985

In this same study, Bridwell relied heavily on character-
istics of the male genitalia. But, since he was unfamiliar with
many of the previously described forms, placement of these
in his scheme was based on imperfect descriptions and he
was not always correct. Cockerell (1942) noted some diffi-
culty in recognizing Bridwell’s groupings. In the main, how-
ever, the classification proposed by Bridwell is sound and
provides a basis on which the present study was constructed.
At the time this revision began, there were 93 species-group
names applied to Ethiopian Region hylaeines.

The first part of this study, in addition to recharacterizing
the previously described genera and subgenera, describes two
new genera and two new subgenera of Hylaeus. These smaller
groups are revised. Regrettably, these small groups appear
to consist of species that are mostly rare or uncommon and
the amount of material available is limited.

Hylaeine bees commonly nest in hollow plant stems, ap-
parently utilizing already excavated sites. Unfortunately, there
have been no studies of the nesting biology of the African
species. Similarly, there are scanty records of the flower vis-
itations for African species. Such data as are available are
cited under each species.

TERMINOLOGY AND MEASUREMENTS
Figures 1-5

Antennal socket diameter (ASD). The maximum diameter,
between the outer margins, at a right angle to the longitudinal
axis of the head in frontal view.

Basal clypeal width (BCW). The distance between the sub-
antennal sutures along the basal margin of the clypeus.

Clypeal length (CL). The median length of the clypeus from
the basal margin to the apical margin; this differs from Hous-
ton’s (1975) measurement, which extends to the level of the

1 . Entomology Section, Natural History Museum of Los Angeles
County, 900 Exposition Boulevard, Los Angeles, California 90007,
U.S.A.

ISSN 0459-81 13

Figures 1-5. Taxonomic characters of Hylaeinae. 1, frontal view of female head; 2, lower face of female; 3, ocellocular area, dorsal view; 4,
posterodorsal view of propodeum; 5, male sternum 7 (lateral teeth stippled). Abbreviations: AOD, antennocular distance; BL, basal lobe;
BTR, basal triangle; CAD, clypeoantennal distance; CW, clypeal width; DL, distal lobe; LAT, lateral carina; MTN, metanotum; OBL, oblique
carina; OCD, ocelloccipital distance; STA, stigmatal area; Tl, tergum 1; TRN, transverse carina. See text (Terminology and Measurements)
for remaining abbreviations.

2 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

lowermost extremity of the clypeus, but is consistent with
my prior usage.

Clvpeo-ocu/ar distance (COD). The minimum distance be-
tween the laterobasal angle of the clypeus and the inner eye
margin.

Frontal shield (FS). For the usually elevated and marginate
area of the face above and between the antennal sockets,
Houston (1975) has proposed “elevations of the interanten-
nal area.” This is cumbersome and I prefer to use the simpler
“frontal shield.” The frontal shield is elevated above the
frons and is distinctly margined and usually widened at about
the midpoint. Houston has noted that the width of the frontal
shield (FSW) at its apex on the frons, when compared to the
diameter of the antennal socket (ASD), is useful as a specific
character.

Flead length (HL). The maximum midline distance be-
tween the occipital margin and the apical margin of the clyp-
eus, in frontal view.

Flead width (HW). The maximum breadth, across the eye,
of the head in frontal view.

Interantenna/ distance (IAD). The minimum distance be-
tween the inner margins of the antennal sockets.

Interocellar distance (IOD). The minimum distance be-
tween the posterior ocelli.

Lower facial width (LFW). The minimum distance be-
tween the eyes at their lower end. This term is utilized in its
relationship to UFW (q.v.) to express degree of convergence
of the inner eye margins: weakly convergent— UFW 1.01-
1.29 x LFW; moderately convergent— UFW 1.30-1.49 x
LFW; strongly convergent— UFW 1.50-1.70 x LFW; very
strongly convergent— UFW more than 1.70 x LFW.

Ocellar diameter (OD). The transverse diameter of the
anterior ocellus.

Ocellocular distance (OOD). The minimum distance be-
tween a posterior ocellus and the inner eye margin.

Pilosity. The distribution of hairs on hylaeine bees, whether
simple or short-plumose, is monotonously uniform, and pi-
losity is not accorded attention here except when it offers
useful differences between species. The following character-
istics apply generally throughout the Ethiopian Region fauna
and will receive no further mention.

The hairs are uniformly whitish; those on the inner surface
of the tarsal segments are usually yellowish, and the ex-
tremely short hairs of the face are often brownish yellow;
females may have a few long dark brown or blackish hairs
on the last tergum and/or sternum.

Very short, simple, suberect to erect hairs are very sparsely
distributed over the surfaces of the head and thorax. Similar,
but slightly longer, appressed to suberect hairs are present
on the abdominal segments and they may be abundant on
the apical terga.

Short, short-plumose, suberect to erect hairs are present
on the face above the level of the antennal sockets, on the
gena (where they become longer toward the mandibular base
and toward the gula), on the side of the thorax, on the side
and disc of the propodeum and on the sides of the abdominal
terga, as well as on the discs of the third and following seg-
ments. They are sparse on all areas except the side and disc

of the propodeum where they are conspicuously more abun-
dant.

Relatively long, erect, short-plumose hairs are present
around the antennal sockets, on the vertex, around the wing
bases, on the side and center of the thorax, and on the apical
terga and sterna. They are most abundant around the anten-
nal sockets and wing bases and on the thoracic venter.

Slightly shorter, mixed barbulate and short-plumose hairs
are present on the mandible, antennal scapes, and the legs.
They are sparse on those segments and are usually suberect
to erect, often moderately curved.

Important specific characters of pilosity are: the presence
of long, erect, short-plumose hairs on the lower face, the
mesoscutum, and the discs of the abdominal terga; the pres-
ence of a transverse fascia of short, appressed, short-plumose
hairs on the pronotal collar; the presence of long, erect, short-
plumose hairs at the juncture of the anterior and dorsal faces
of the first tergum and/or at the side of that segment; the
presence of fasciae, entire or not, of short, appressed, short-
-plumose hairs preapically on any of the abdominal terga.
Although other differences between species do exist, they are
often too subtle to be useful and are, therefore, ignored here.

Pronotal lobes. Houston (1975) proposed to call the dorsal
and posterior lobes of the pronotum the “pronotal collar”
and “pronotal tubercles,” respectively. Since Michener (1965)
has already utilized the former following a tradition estab-
lished by earlier workers, the usage is continued here, as it
has been in my earlier papers. In place of “pronotal tuber-
cles,” I still prefer the older term “pronotal lobes,” for they
are lobes and not tubercles.

Propodeum. The hylaeine propodeum is useful in deter-
mining group relationships, and it is convenient to refer to
specific subdivisions in the descriptions. In dorsal view, the
most obvious feature is the more or less horizontal basal or
dorsal face; a pair of sutures extends from the antero-lateral
margin, converging toward the postero-median margin of the
basal area. The resultant somewhat triangular area is the
basal triangle, the apex of which is situated on the posterior,
more or less vertical surface, here called the propodeal disc;
the disc is divided by a median groove, the sides of which
are continuous with the margins of the basal triangle. The
lateral margin of the disc, at its juncture with the side of the
propodeum, is often marked by a low carina, the lateral
carina, which may extend forward to the anterior margin of
the propodeum, but usually does not. At the point where the
lateral carina reaches the basal face there may be another low
carina which extends obliquely mesad to join the side of the
triangle; this is the oblique carina. The area mesally bounded
by the margin of the triangle, posteriorly by the oblique carina
and laterally by the lateral carina (or its imaginary extension)
is the stigmata/ area (after Benoist, 1959) and is equivalent
to the “lateral area” of Dathe (1980).

Punctation. I prefer to use puncture in preference to Hous-
ton’s (1975) “pit”; the former is long established and con-
sistent with general terminology used in apoid systematics.
Houston illustrated his various terms (minute, fine, small,
medium, large, coarse) to express relative size of punctures,
but I prefer more absolute definitions. Puncture diameters

Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees 3

are measured, by means of an ocular micrometer, at 120x,
and the following terminology applies to various puncture
diameters:

minute 0.010-0.019 mm

fine 0.020-0.035 mm

moderate 0.036-0.055 mm

coarse 0.056-0.070 mm

very coarse over 0.070 mm

Since punctures are rarely of one size on a given segment
or stipulated area, they may be described as “fine to mod-
erate” (puncture diameter varying between 0.020 and 0.055
mm), though usually a more limited size range, such as “mod-
erate” prevails.

The relative density of the punctation on a given segment
is often different in closely related and otherwise similar
species. The commonly applied terms such as close, dense,
sparse, etc., are usually not defined and subject to consid-
erable latitude of interpretation. I have attempted to stan-
dardize my terminology for ease in comparison and, perhaps,
encourage some degree of accepted usage. Because the in-
terspaces within an indicated area are somewhat variable in
extent, the stated condition in descriptions is that which is
prevalent:

Contiguous punctures are so close that they are often de-
formed; their interspaces are greatly compressed and sharp
edged.

Subcontiguous punctures are separated by more or less flat-
topped interspaces up to about 0.25 puncture diameters; at
its lower extreme this merges into contiguous.

Dense punctures are separated by more or less flat-topped
interspaces varying from about 0.30 to about 0.70 puncture
diameters; most commonly about 0.50 puncture diameters.

Close punctures are separated by more or less flat-topped
interspaces varying from about 0.70 to about 1 .50 puncture
diameters.

Sparse punctures are separated by distances from 2.00 to
about 3.00 puncture diameters.

Scattered punctures are separated by very irregular inter-
spaces, from about 3.00 to as much as 6.00 puncture di-
ameters, often with extensive areas devoid of punctation.

Since size and density of punctation are often not uniform
on all areas of a given segment, the following comments are
necessary. The description of clypeal punctation is derived
from the basal one-third of that segment; punctures usually
are somewhat coarser toward the apex and often are sparser
along the midline. Genal punctation is finest on the upper
one-third and becomes gradually coarser, closer and more
distinct toward the base of the mandible.

The mesoscutal punctation is described from the area be-
tween the midline and the parapsidal line at the level of the
tegula; punctures become finer and, often closer, anterior to
this area; often coarser and somewhat sparser in the postero-
median area, but very fine and dense along the mesoscutal-
scutellar suture. Scutellar and metanotal punctation are
described from the mesal one-third of each segment. The
middle of the mesopleural disc is the standard for that seg-
ment as is also true for the side of the propodeum.

Scape length (SL). The standard measurement, exclusive
of the basal condyle.

Scape width (SW). The maximum width of the scape.

Sculpture. Except for the obvious differences related to
punctation, I have not devoted much attention to superficial
texture. My main reason is simply that the superficial texture
varies considerably within a species, and the differences be-
tween closely related species are often so subtle that descrip-
tions are useless and/or largely subjective. So, I use “tessel-
late” generally to describe the surface: “lightly tessellate”
corresponds approximately to Houston’s “lineo-reticulate”
and “closely tessellate” is approximately equivalent to his
“pit-reticulate”; lineolate is the same as his “transversely
lineo-reticulate.”

Total length (TL). This is the least satisfactory of mea-
surements used here; it is certainly the least exact. The meth-
od used here differs from the conventional ones, but seems
less subject to the vagaries resulting from wide variations in
the death posture of the specimen or its final, mounted con-
dition. The TL is derived by adding the following: HL +
thoracic length (in dorsal view, along the midline, from the
anterior margin of the pronotal collar to the posterior margin
of the dorsal or basal face of the propodeum) + length of
tergum 1 (dorsal view along the midline, with the summit
of the basal face just occluding the basal attachment) + length
of tergum 2 (along midline, from gradulus to apical margin).

Upper facial width (UFW). The minimum distance be-
tween the eyes above, at about the level of the anterior ocel-
lus, or somewhat below, but not at the point of greatest width
as Houston (1975) has it. The usage here is consistent with
that in my earlier papers.

Wing length (WL). The length of the anterior wing, from
the tegular margin to the wing apex.

DESCRIPTIONS

The descriptions of genera and subgenera are divided into
three sections. First is a Diagnosis, a brief statement of the
outstanding features of the taxon. This is followed by the
Description. The Description consists of numbered state-
ments, and they are directly comparable at the appropriate
(i.e., generic or subgeneric) level. Concluding is another state-
ment, supplemental to the Description, of additional char-
acteristics of taxonomic interest.

The species descriptions are more detailed than those of
the genera and subgenera. Although the statements are not
numbered, a uniform descriptive format is used, so descrip-
tions of species are comparable.

Previously described species are usually not redescribed.
In most instances the characteristics noted in keys and dis-
cussions should be ample. All new species are fully described.

ABBREVIATIONS OF MUSEUMS

AMNH American Museum of Natural History, New York,
New York, U.S.A.

BMNH British Museum (Natural History), London, U.K.

4 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

CAS California Academy of Sciences, San Francisco,
California, U.S.A.

CORN Cornell University, Ithaca, New York, U.S.A.

DEI Deutsches Entomologisches I nstitiit, Eberswalde
bei Berlin, D.D.R.

GEMB Faculte des Sciences Agronomiques de Fetat,
Gembloux, Belgium

LACM Natural History Museum of Los Angeles County,
Los Angeles, California, U.S.A.

MCZ Museum of Comparative Zoology, Cambridge,
Massachusetts, U.S.A.

MNHN Museum National d’Histoire Naturelle, Paris,
France

MNHU Museum fur Naturkunde der Humboldt-Univer-
sitat, Berlin, D.D.R.

MRAC Musee Royal de FAfrique Central, Tervuren, Bel-
gium

PRET National Collection of Insects, Pretoria, South Af-
rica

SAM South African Museum, Cape Town, South Africa
[In 1 98 1 the Hymenoptera collections of the Na-
tional Museum of Rhodesia (now Zimbabwe),
Bulawayo, were transferred to the South African
Museum, Cape Town. Material borrowed from
the Rhodesian National Museum is, therefore,
included herein under SAM.]

UKAN University of Kansas, Lawrence, Kansas, U.S.A.
USNM National Museum of Natural History, Washing-
ton, D.C., U.S.A.

LIST OF EXCLUDED SPECIES

The following species are excluded for reasons stated under
each name.

Prosopis albonasata Strand, 1912. Described from a single
male from “Kapland.” The type has been examined, and I
conclude that the locality may be spurious. This is a synonym
of the Palaearctic species, Hylaeus ( Prosopis ) signatus (Pan-
zer) (NEW SYNONYMY).

Prosopis gracilis Bingham, 1903. Described from a female
presumably from Durban, Natal, in BMNH. According to
Michener (1975), a synonym of Braunsapis facialis (Ger-
staecker) (Anthophoridae).

Prosopis pernix Bingham, 1903. Described from a female
from Port Natal, Natal. According to Michener (1975), this
is a valid species in Allodape (Anthophoridae).

Prosopis quadrilineata Cameron, 1905. Described from a
female from Grahamstown, Brak Kloof, in BMNH. Mich-
ener (1975) treats this as a valid species in Allodape.

Prosopis 5-lineata Cameron, 1905. Described from a fe-
male from Stellenbosch, Cape Province, in BMNH. Accord-
ing to Michener (1975), this is a synonym of Allodape pic-
tifrons F. Smith.

Prosopis sandaracta Bingham, 1903. Described from a fe-
male from Durban, Natal, in BMNH. Friese (1909) stated
that this is a synonym of Hylaeus purpurisata (Vachal) of
Algeria, but there is no evidence that he actually saw the
type. According to Cockerell (1934), this is a synonym of

Allodapula variegata (F. Smith) (Anthophoridae) with which
Michener (1975) concurs.

KEY TO GENERA AND SUBGENERA OF
ETHIOPIAN REGION

1. Integument of both sexes black, red or both, never

strongly metallic; metatibia of female without raised gla-
brous area at base; male genitalia with well-developed,
cup-like base 2

- Integument of both sexes metallic blue to blue-green;

metatibia of female with raised glabrous area at base;
male genitalia with gonobase reduced to a basal ring (Fig.
61) Calloprosopis, new genus

2. Supraclypeal area elevated between antennal sockets and

laterally marginate; propodeum with defined basal area
and usually coarsely rugose or roughened or sharply
punctate, at least in part 3

- Supraclypeal area gently sloping from midline to anten-
nal sockets, not laterally marginate; propodeum smooth,
densely tessellate, without defined basal area; entire body
densely tessellate, without conspicuous punctures ....

Psilylaeus, new genus

3. Apical margin of mandible transverse or oblique, bi- or

tridentate, mandible short and broad (Figs. 12-15) ( Hy-
laeus) 4

- Apical margin of mandible acute, without distinct teeth,

mandible elongate and slender (Figs. 16-17)

Not hylaeus Bridweli

4. Integument variously punctate; scutellum and metano-
tum without lateral spines; occipital carina often absent;
mesepisternum not sharply carinate, but sometimes with
obscure ridge at juncture of lateral and anterior faces

5

- Integument very coarsely punctate; scutellum and meta-

notum each usually with a pair of spines; occipital carina
present, sharp; mesepisternum sharply carinate at junc-
ture of anterior and lateral faces

subg. Metylaeus Bridweli

5. Male, antenna 1 3-segmented, gaster with 7 visible seg-
ments 6

- Female, antenna 1 2-segmented, gaster with 6 visible seg-
ments 9

6. Gonocoxite abruptly narrowed over apical one-third or

more (Figs. 31, 36, 40, 45) 7

- Gonocoxite terminating bluntly at level of apex of ae-

deagus (Figs. 21, 50) 8

7. Face and first tergum coarsely, closely punctate; lateral

lobes of sternum 7 thin, strap-like, very short (Fig. 29);
apex of gonocoxite without barbulate setae

Alfkenylaeus, new subgenus

- Face finely punctate, punctures obscured by tessellation;

lateral lobes of sternum 7 bifurcate, basal lobule with a
few coarse teeth on margin (Figs. 5, 6); apex of gono-
coxite with two or three barbulate setae

subg. Deranchylaeus Bridweli (part)

8. All terga with abundant erect, fine, white hairs, these
becoming longer and denser caudad (Fig. 18); clypeus

Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees 5

black or with median yellow stripe which tapers toward
apex, but not reaching apical margin; terga 1 and 2
coarsely, closely punctate; basal lobule of sternum 7 short,
without lateral teeth Cornylaeus, new subgenus

- Discs of terga 1-3 usually with few or no erect hairs (if

any present, they are short, separated by their own lengths
or more and often dark); clypeal marks variable, but
rarely as described above, clypeus usually entirely pale;
terga 1 and 2 often without evident punctures; basal
lobule of sternum 7 often elongate and always laterally
dentate subg. Deranchylaeus Bridwell (part)

9. Mandibular apex bi- or tridentate, dorsal border usually
not flattened and expanded, but if flattened, conspicu-
ously punctate and only slightly shiny and pronotal col-
lar sharply marginate in middle; clypeal marks various,
often greatly reduced or absent; tergal fasciae and punc-
tation various 10

- Mandibular apex bidentate, dorsal border flattened and

expanded, shiny and impunctate (Fig. 12); lateral face
mark full, clypeus black or with median stripe, pronotal
stripe complete; first tergum conspicuously punctate; ter-
ga 1 and 2 with apical pubescent fasciae, broadly inter-
rupted in middle; pronotal collar not marginate in mid-
dle Cornylaeus, new subgenus

10. Clypeus flat preapically without conspicuous transverse

or quadrate impression 11

- Clypeus with pronounced median, preapical quadrate or

somewhat transverse depression

subg. Deranchylaeus Bridwell (part)

1 1 . Clypeus coarsely, closely punctate; pronotum sharply

marginate in middle; lateral face mark complete, but not
extending above level of antennal socket; legs and clyp-
eus often ferruginous Alfkenylaeus, new subgenus

- Clypeus usually finely and sparsely punctate; pronotum

usually rounded in middle; if clypeus and pronotum as
above, then lateral face mark absent or reduced to nar-
row stripe along eye margin; legs and clypeus not fer-
ruginous, or, if ferruginous, then clypeus finely punctate
and lateral face mark absent

subg. Deranchylaeus Bridwell (part)

Hylaeus Fabricius

Hylaeus Fabricius, 1793:302. Type-species: Apis annulata
Linnaeus, 1 758; designation ofLatreille, 1810.

DIAGNOSIS

Body mostly dull black, with or without pale marks on head
and thorax; frontal shield present; mandibles stout, bi- or
tridentate; at least head and thorax usually conspicuously
punctate.

DESCRIPTION

( 1 ) Mandible stout, bidentate in male, bi- or tridentate in
female. (2) Labral tubercle, when present, small, midbasal.

often depressed along center. (3) Tentorial pit usually at mid-
point of clypeal length. (4) First flagellar segment, and often
second as well, short and transverse. (5) Frontal shield pres-
ent and sharply margined. (6) Lateral carina of propodeum
usually present, oblique carina usually absent in our fauna.
(7) Sulcus of first tergum long or short. (8) Gradulusof second
tergum gently arched, usually evanescent laterad. (9) Third
tergum of male without sublateral pubescent fovea. (10) Male
sternum 7 normally with distinct basal and apical lobules.

(11) Male sternum 8 usually with distal process long and
slender; dorsal tubercle usually near base of distal process.

(12) Male gonocoxite usually broad and not extending much
beyond penis valves, but may be elongate and slender.

DISCUSSION

Hylaeus is a virtually cosmopolitan genus with many species
arrayed within numerous subgenera. The European species
have been recently reviewed by Dathe (1980), and many of
the Nearctic species by Snelling (1966a-c, 1968, 1970). The
hylaeines of Australia are presently being revised by Houston
(1975, 1981). The extensive Neotropical and Asian faunas
are essentially unstudied.

The above description is drawn entirely from the species
of the Ethiopian Region and will not apply, in all particulars,
to Hylaeus from other Regions. Species are mostly small
black bees with limited whitish or yellowish marks on the
head and thorax, especially in the males. A few species have
limited red marks, particularly on the clypeus and legs.

After removing some species to Not hylaeus and Metylaeus,
Bridwell (1919) placed all remaining Ethiopian hylaeines in
his subgenus Deranchylaeus. Many of those included were
known to Bridwell only from their original descriptions. It
is not surprising, therefore, to find that some will not fit
within Bridwell’s scheme. For these species, new subgeneric
or generic names are proposed. At the same time, Bridwell’s
genus Metylaeus is treated as a subgenus of Hylaeus.

Most species still remain in Deranchylaeus, a large and
diverse subgenus, apparently restricted to the Ethiopian Re-
gion. Deranchylaeus may be derived from the large Holarctic
subgenus Prosopis, or at least from a similar stock. However,
until the taxa within Hylaeus can be studied on a worldwide
basis, this must be presumption only.

Subgenus Deranchylaeus Bridwell

Hylaeus subg. Deranchylaeus Bridwell, 1919:1 36-137. Type-
species: Prosopis curvicarinata Cameron, 1905; original
designation.

DIAGNOSIS

Mandibles short, broad, bi- or tridentate at apex; preoccipital
carina absent; mesepistemum not carinate between anterior
and lateral faces; scutellum and metanotum unmodified; male

Figures 6-11. Sterna 7 and 8, genital capsule (right half dorsal, left half ventral views) of: 6-8, H. (Deranchylaeus) sp., scale line = 0.25 mm;
9-11, H. ( Nothylaeus ) heraldicus, scale line = 0.50 mm.

6 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

Contributions in Science, Number 361

SneMing: Ethiopian Hyiaeine Bees 7

sternum 7 with lateral margin of basal lobule dentate; male
gonocoxite stout, ending at about level of penis valves.

DESCRIPTION

( 1 ) Mandible broad, usually bidentate, but tridentate in some
females. (2) Clypeus flat or with distinct preapical depression;
punctures minute to fine, usually sparse but may be close.
(3) Preoccipital carina or ridge absent. (4) Pronotal collar
usually rounded on dorsal surface, rarely somewhat margin-
ate. (5) Mesepistemum without carina between anterior and
lateral faces. (6) Lateral carina of propodeum usually present,
oblique carina usually absent. (7) Sulcus of tergum 1 one-
third or less as long as basal face. Tergum 2 with punctiform
lateral fovea. Male sterna 2 and 3 usually not tuberculate.
(10) Male sternum 7 with teeth along lateral margin of basal
lobule. (11) Male gonocoxite broad, blunt, extending little,
if any, beyond level of apex of penis valves.

Dorsal border of mandible usually not flattened, but if so,
expanded area is densely punctate and only slightly shiny;
scutellum and metanotum simple.

DISCUSSION

This is the largest group of hylaeines in the Ethiopian Region.
Many of the species appear to be common and widespread.
These wide-ranging species are subject to considerable vari-
ation in the intensity and distribution of pale face marks,
with many trivial forms named on the basis of slight color
differences. There are 10 clearly recognizable species groups.

LIST OF INCLUDED SPECIES NAMES

absonulus Cockerell, 1936a
abjunctus Cockerell, 1936a
alfkeni ( Friese, 1913)
atriceps (Friese, 1911)
bequaertianus Bridwell, 1919
capicola (Alfken, 1914)
clavigerus Cockerell, 1936b
corpana (Wamcke, 1972)
curvicarinatus (Cameron, 1905)
dominae Cockerell, 1936a
dregei (Strand, 1912)
extensicornis Cockerell, 1936a
flaviscutum (Alfken, 1914)
gabonica (Vachal, 1899)
graafi Cockerell, 1936a
haygoodi Bridwell, 1919
immarginatus (Alfken, 1914)
kasindensis Cockerell, 1936a
krebsianus (Strand, 1912)
lemuriae (Benoist, 1946)*

* Known only from Madagascar.

leucolippa ( Friese, 1913)
lightfooti Bridwell, 1919
lineaticeps ( Friese, 1913)
longula (Friese, 1913)
major (Strand, 1912)
malagassa (Benoist, 1946)*
melanosomus Cockerell, 1920
ogilviei Cockerell, 1936
perater Cockerell, 1936a
perdensus Cockerell, 1936a
promontorii Meade-Waldo, 1923
punctifrons Cockerell, 1936a
punctiferus Cockerell, 1936a
reditus Cockerell, 1936a
rhodognathus Cockerell, 1936a
robertiana (Cameron, 1906)
rugipunctus (Alfken, 1914)
sanctus Cockerell, 1936a
simplex (Bingham, 1912)
simplior Meade-Waldo, 1923
simulans Cockerell, 1942
stictifrons (Cockerell, 1936b)
sublucens Cockerell, 1936a
subreditus Cockerell, 1 942
tenuis (Alfken, 1914)
tinctulus Cockerell, 1932
varians Cockerell, 1936a
vau Cockerell, 1936a
xanthostoma (Alfken, 1914)

Cornylaeus, new subgenus

Type-species: Prosopis aterrima Friese, 1911.

DIAGNOSIS

Mandible bidentate, that of female with upper margin flat-
tened, expanded, shiny and impunctate; mesepistemum not
carinate between anterior and lateral faces; scutellum and
metanotum simple; macula of pronotal collar complete; male
with abundant fully erect hairs on discs of abdominal seg-
ments; male sternum 7 without teeth on lateral margin of
basal lobule; male gonocoxite stout, ending at about level of
apex of aedeagus.

DESCRIPTION

( 1 ) Mandible bidentate at apex, upper border flattened and
expanded in female, expanded portion shiny and impunctate.

(2) Clypeus flat, without preapical impression. (3) Preoccip-
ital ridge or carina absent. (4) Pronotal collar rounded above.
(5) Mesepistemum without carina between anterior and lat-
eral faces. (6) Oblique carina absent, lateral carina present
in its lower half only. (7) Sulcus of tergum 1 about one-third
as long as basai face. (8) Tergum 2 with punctiform lateral
fovea. (9) Male sternum 3 conspicuously tuberculate, or not.

Figures 12-17. Frontal view of head of female and male of: 12-13, H. (Cornylaeus) aterrimus: 14-15, H. ( Cornylaeus ) proteae. ; 16-17,
Nothylaeus heraldicus. Scale line = 1.00 mm. Figures by R.A. DeNicola.

8 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees 9

(10) Male sternum 7 without teeth along lateral margin of
basal lobule. (11) Male gonocoxite broad, blunt, ending slightly
beyond apex of penis valves.

Male labrum without median tubercle; IAD about 0.66 x
COD; subantennal sutures about as long as ASD; upper end
of female facial fovea slightly separated from inner eye mar-
gin; first flagellar segment slightly broader than long, longer
than second segment; male mesepistemum with longitudinal
tubercle below.

DISCUSSION

This subgenus is proposed for two apparently uncommon,
rather large and robust forms. Although Bridweli (1919) in-
cluded the type species in his subgenus Deranchylaeus, there
is no evidence that he actually saw any specimens. Males are
easily recognizable by the presence of abundant long erect
hairs on the discs of the abdominal segments. In most males,
too, there are conspicuous tubercles on the third sternum.

The males of at least one species, H. aterrimus, are poly-
morphic, exhibiting a wide range of variation in the devel-
opment of the tubercles of sternum three. In this species,
tubercles may also be present on the third tergum. In general,
degree of development of the tubercles is correlated with
body size, but there is no consistency. So far as known, males
of H. proteae always possess a large, somewhat asymmetrical
process on the third sternum, but too few specimens have
been seen for there to be any certainty about this.

Females of Cornylaeus are less easily recognized, but the
mandibular structure is unlike that of any Deranchylaeus. In
addition, the basal face of the first tergum has numerous fully
erect white hairs, usually long and conspicuous. These hairs
are subject to wear and are sometimes absent. A few De-
ranchylaeus do possess hairs on the basal face of the first
tergum but do not have the mandibular structure character-
istic of Cornylaeus.

ETYMOLOGY

The subgeneric name is derived by combining the Latin word
for horn, or tubercle ( cornus ), with the name Hylaeus.

SYNONYMIC LIST OF SPECIES

aterrimus (Friese)

=quinquedentata Friese
^pondonis Cockerell, NEW SYNONYMY
proteae Cockerell

KEY TO SPECIES OF CORNYLAEUS

1 . Female, antenna 1 2-segmented 2

- Male, antenna 13-segmented 3

2. Clypeus with median stripe; supraclypeal area maculate

aterrimus (Friese)

- Clypeus and supraclypeal area black . . proteae Cockerell

3. Clypeus with longitudinal stripe; metatibia with basal pale

mark; tergum 3 often with lateral tubercles; sternum 3
tuberculate or not; scape longer than broad

aterrimus (Friese)

- Clypeus black or with a minute preapical median spot;
metatibia wholly dark; tergum 3 always without lateral
tubercles; sternum 3 with a large median swelling; scape
rotund, as broad as long proteae (Cockerell)

Hylaeus ( Cornylaeus ) aterrimus (Friese)

Figures 12-13, 18-21

Prosopis aterrima Friese, 1 9 1 1 : 1 20. 3, 9. SOUTH AFRICA:
Shilouvane, N. Transvaal, Feb. (9), Oct. (3) (MNHU) [ex-
amined].

Prosopis quinquedentata Friese, 1911:132. <5. SOUTH AF-
RICA: Shilouvane, N. Transvaal (Junod) (MNHU) [ex-
amined].

Hylaeus pondonis Cockerell, 1942:10. <3. SOUTH AFRICA:
Port St. John, Pondoland, Oct. 1923 (R E. Turner) (BMNH)
[examined], NEW SYNONYMY.

Three cotypes are available from MNHU. Of these, the male,
which agrees with Friese’s original description, is selected as
lectotype. Of the two females, the smaller is designated al-
lolectotype and the larger paralectotype; all specimens are in
MNHU.

Alfken (1914) first recognized that H. quinquedentata was
a junior synonym of H. aterrima. I have examined the type
and concur. Cockerell’s H. pondonis is based on a male lack-
ing tubercles on the third tergum. It falls well within the
range of variation of H. aterrimus. as I understand the species.

MATERIAL EXAMINED

CONGO REPUBLIC: 286, 10 mi. S Kapona, 1570 m elev.,
13 Jan. 1958 (E.S. Ross & R E. Leech; CAS). ZIMBABWE:
19, 13, Salisbury, no date (D. Dodds; SAM, AMNH); 13,
Burnside, Bulawayo, 28 Sept. 1952 (no name; SAM); 288,
Bulawayo, 24 Apr. 1916 (no name; SAM); 19, Bulawayo,
Apr. 1916 (no name; SAM); 19, Bulawayo, 29 Apr. 1916 (no
name; SAM); 13, Bulawayo, 6 June 1925 (R.H.R. Stevenson;
SAM); 13, Bulawayo, 7 Sept. 1954 (no name; SAM); 19, Hope
Fountain, 7 May 1916 (no name; SAM); 13, Salisbury, Mar.
1906 (G.A.K. Marshall; BMNH); 233, Umtali Heights, 1420
m elev., 13 Mar. 1958 (E.S. Ross & R.E. Leech; CAS).
SOUTH AFRICA: 19, Port St. John, Pondoland, 1-17 Mar.
1924 (R.E. Turner; BMNH); 13, same locality and collector,
Jan. 1924 (BMNH); 19, same locality and collector, Oct. 1923
(BMNH); 233, Hilton, Natal, 2 Aug. 1966 (J.S. Taylor;
USNM), on Protaea; 233, Hellabella, 2200 ft. elev., 12 mi.
SW Richmond, 13 Jan. 1967 (C.D. Michener & D.J. Broth-
ers; UKAN); 13, 299, Shilouvane, Transvaal, no date (Junod;
MNHU, cotypes of P. aterrima ); 13, same data (MNHU,
type of P. quinquedentata ); 13, Wolkberg, Transvaal, 1 3 Apr.
1974 (R.H. Watmough;BRET)\ 19, Long Tom Pass, 25°07'S,
30°35'E, Transvaal, Jan. 1977 (E.F. Whitehead; PRET),
“yellow bowl trap.”

Hylaeus ( Cornylaeus ) proteae Cockerell

Figures 14-15, 22-26

Hylaeus proteae Cockerell, 1942:1 1-12. 3. SOUTH AFRI-

10 Contributions in Science, Number 361

Sneiling: Ethiopian Hylaeine Bees

(

Figures 18-21. Male, H. ( Cornylaeus ) aterrimus : 18, lateral habitus (scale line = 2.00 mm); 19-21, sterna 7 and 8, genitalic capsule (scale
line = 0.50 mm). Figure 18 by R.A. DeNicola.

Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees 1 1

Figures 22-26. Male, H. ( Cornylaeus ) proteae: 22, abdomen, lateral view; 23, sternum 3, ventral view; 24-26, sterna 7 and 8 genitalic capsule
(scale line = 0.50 mm, 24-26 only). Figures 22 and 23 by R.A. DeNicola.

CA: Port St. John, Pondoland, Oct. 1923 (R.E. Turner)
(BMNH) [examined].

The male of H. proteae is easily recognized by the combi-
nation of black clypeus, externally globose scape, densely
pubescent abdomen and enormous tubercle on the third ster-
num. In the female the cutting margin of the mandible is
oblique behind the second tooth, the blade is broad, the

clypeus is black, and the pronotal collar has a continuous
yellow band.

MATERIAL EXAMINED

ZIMBABWE: 1 2, Vumba Mts., 27 Feb. 1 938 (no name; SAM);
222, same locality, 12 Aug. 1956 (no name; SAM). SOUTH
AFRICA: 222, 433, Port St. John, Pondoland, Oct. 1 923 (R.E.

12 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

Turner; BMNH, SAM, including 2 cotype <3<3), on Protaea;
12, Magaliesberg, Tonguani Kloof, Transvaal, 2 Feb. 1975
(P.H. Watmough; PRET), in “open grassland.”

Alfkenylaeus, new subgenus

Type-species: Hylaeus namaquensis Cockerell, 1942.

DIAGNOSIS

Male with gonocoxite abruptly narrowed over distal one-
third or more; seventh sternum with apical lobule virtually
absent, basal lobule strap-like and without teeth. Female
mandible weakly tridentate; gradulus of second tergum weak-
ly curved, deflected at side; pregradulus of third sternum
elevated and extended caudad in middle. In both sexes, head,
thorax (except propodeum in one species) and first two terga
coarsely and closely punctate; pronotal collar carinate.

DESCRIPTION

(1) Male mandible bidentate at apex, that of female weakly
tridentate. (2) Clypeus gently arched, without preapical
impression. (3) Occipital margin acute, but preoccipital ridge
absent. (4) Pronotal collar carinate on its dorsum. (5) Mes-
epistemum without carina between anterior and lateral faces.
(6) Oblique and lateral propodeal carinae absent. (7) Sulcus
of tergum 1 broad at base, at least half as long as basal face,
depressed and marginate. (8) Tergum 2 with broadly oval
lateral fovea; gradulus gently bowed in middle, weakly de-
flected at side. (9) Male sternum 3 with or without transverse
swelling. (10) Male sternum 7 with distal lobules reduced or
absent, basal lobules strap-like and without teeth (Figs. 29,
34, 38). Male gonocoxite abruptly narrowed in distal one-
third or more, ending much beyond apex of penis valves
(Figs. 31, 36, 40), usually protruding from genital opening
in dry specimens.

IAD subequal to (H. namaquensis) or much less than (H.
acariphorus and H. psaenythioides) COD; frontal shield high,
narrow (more so in males), sharply marginate and reflexed
at sides; female facial fovea ending nearer eye than ocellus;
first two flagellar segments transverse, subequal in length or
second slightly longer; pregradulus of tergum 3 elevated and
narrowly, triangularly extended distad in middle; distal pro-
cess of male sternum 8 elongate, narrow, and slightly broad-
ened at apex (Figs. 30, 35, 39).

ETYMOLOGY

This subgenus is dedicated to J.D. Alfken whose early work
on Hylaeus has contributed greatly to our understanding of
the Old World forms.

DISCUSSION

This subgenus is proposed to accommodate a few coarsely
punctate species of distinctive habitus. The unusually large
basal sulcus of the first tergum is apparently a modification
to accommodate the mites often found on these bees. The

elevated pregradular area of the third segment may be sim-
ilarly adaptive.

The following key to species of Alfkenylaeus includes both
sexes of H. arnoldi (Friese), although this bee does not belong
to this subgenus. In the key to genera and subgenera, how-
ever, H. arnoldi will come out with the Alfkenylaeus species
and so it seems most convenient to include the species here;
H. arnoldi is discussed following treatment of the species of
Alfkenylaeus.

SYNONYMIC LIST OF SPECIES

acariphorus. new species
infulatus, new species
namaquensis Cockerell
psaenythioides, new species

KEY TO SPECIES OF ALFKENYLAEUS

1. Female, antenna 12-segmented 2

- Male, antenna 13-segmented 6

2. Apical pro tarsal segment broadest at apex, evenly nar-
rowed toward base; bristles of protarsus slightly or not at
all flattened, their apices acuminate; terga 1-2 coarsely,

closely punctate 3

- Apical protarsal segment narrowest basad, broadened to
basal one-third, evenly narrowed apicad; protarsal bristles
strongly flattened, apices bluntly rounded; second tergum
either without evident punctures or punctures fine and

obscure arnoldi (Alfken)

3. Clypeus entirely black; legs brownish, metatibia with bas-
al one-third more or less yellowish; terga 2-5 with con-
spicuous preapical pubescent fasciae 4

- Clypeus largely ferruginous; legs entirely ferruginous; ter-

ga 3-5 with short, inconspicuous hairs, a little denser on
side of first and second segments

acariphorus, new species

4. Punctures along middle of clypeus fine to moderate, sep-

arated by one-half, or more, puncture diameters; pronotal
collar maculate; hairs on basal face of tergum 1 long, fully
erect, and continuous across summit

namaquensis Cockerell

- Punctures along middle of clypeus coarse and contiguous
to subcontiguous; pronotal collar immaculate; hairs of
basal face of tergum 1 subappressed and limited to margin
of sulcus, none across summit . . infulatus, new species

5. Clypeus entirely pale or pale with apical area ferruginous;

lateral face mark filling most of space between eye and
clypeus; thorax often dull, punctural interspaces densely
tessellate 6

- Clypeus black on basal two-thirds, apical one-third fer-

ruginous; transverse stripe between antennal sockets and
clypeal base yellowish; thoracic interspaces shiny, pol-
ished on dorsum psaenythioides, new species

6. Sternum 3 flat, without mediobasal glabrous swelling; base

of metatibia and entire metatarsus whitish 7

- Sternum 3 with low, semicircular glabrous swelling at
base; legs wholly ferruginous . . acariphorus. new species

Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees 13

7. Lateral face mark ending at level of upper margin of an-
tennal sockets; scape maculate beneath; third and follow-
ing terga without preapical pubescent fascia

arnoldi (Alfken)

- Lateral face marking ending midway between antennal
sockets and top of eye; scape immaculate beneath; tergum

3, at least, with preapical pubescent fascia

namaquensis Cockerell

Hylaeus ( Alfkenylaeus ) acariphorus, new species

Figures 27-3 1

DIAGNOSIS

Male: Sternum 3 with large, flat, glabrous swelling in middle
of base; legs wholly ferruginous; clypeus yellow and ferru-
ginous; interspaces of frons and thoracic dorsum slightly shiny,
distinctly tessellate. Female: Terga 2 and 3 with pubescent
fasciae, interrupted in middle; interspaces of frons and tho-
racic dorsum dull, densely tessellate; clypeus red and black,
legs wholly red.

DESCRIPTION

MALE (HOLOTYPE). Measurements. HL 1 .74; HW 1 .97;
SL 0.74; WL 4.9; TL 7.3 mm.

Head. Broad, HW 1 . 1 x HL; scape moderately long, 2.3 x
longer than wide, SL 0.42 x HL. Eyes strongly convergent
below, UFW 1.53 x LFW. Clypeus slightly longer than wide
at apex, sides regularly divergent to maximum width, BCW
0.48 x CW; BCW:COD:CAD:ASD:IAD = 10:10:8:6:5. OD:
IOD:OOD = 5:12:7.5. Frontal shield very narrow between
antennal sockets, sides reflexed, transparent. First flagellar
segment shorter than either pedicel or second flagellar seg-
ment. Entire face coarsely punctate, punctures about 0.06
mm diam.; interspaces in maculate areas slightly shiny and
lightly tessellate, in immaculate areas dull, closely tessellate.

Thorax. Pronotal collar sharply carinate in front, carina
extending laterad to base of pronotal lobe; humeral ridge
present. Mesoscutum 1.35 x wider than long. Scutellum flat,
about 0.37 x length of scutum. Metanotum flat, sloping,
about half as long as scutellum. Basal face of propodeum
gently curved into declivitous face; basal triangle sharply
marginate. Sides of pronotal collar coarsely, closely punctate;
scutum, scutellum, pleura coarsely, almost contiguously
punctate; sides and stigmatal area of propodeum closely
punctate, punctures about one-half size of mesopleural punc-
tures; basal triangle rugosoreticulate. Integument barely shiny,
interpunctural spaces lightly to densely tessellate.

Abdomen. Tergum 1, from above, about 1.4x wider than
long, basal sulcus sharply margined, deep, over half length
of basal face, apical band broad, a little depressed at sides;
apical band about twice as wide on second segment, strongly
depressed at sides; first two terga coarsely, closely punctate;
sternum 3 with large, flat, semicircular, mediobasal, glabrous
tumescence occupying about ‘A of segment; sternum 7 with-
out hairs on apical lobes; sternum 8 abruptly broadened
preapically, apex angulate; gonocoxite evenly narrowed, apex
slightly broadened.

Pilosity. Clypeus with numerous short, erect hairs; sides
and front of face with hairs conspicuously longer, weakly
plumose, especially around antennal insertions; genal hairs
sparse, mostly reclinate. Thoracic dorsum with short, sparse
erect hairs, except around wing bases and sides of scutellum
and metanotum where they are much longer; pleura with
hairs short, sparse above, becoming longer ventrad; propo-
deal hairs sparse, moderately long. Tergum 1 with a few short
simple hairs near base and with fine, appressed plumose hairs
on sides of apical depression; tergum 2 with sparse simple,
erect hairs and dense, plumose, appressed hairs in apical
depression, narrowly interrupted in middle; remaining terga
with sparse simple hairs of variable length, appressed to fully
erect; sterna with scattered, erect, weakly plumose hairs, more
abundant caudad.

Color. Black; mandible, lower sides of face, apical third of
clypeus (except median stripe), scape, flagellum, and legs light
ferruginous; minute basal spot on mandible, basal third of
clypeus and broad median intrusion into apical two-thirds,
supraclypeal area, sides of face to slightly above level of
antennal sockets, all light yellowish. Upper side of scape and
flagellum brownish. Tegula testaceous. Wings clear, veins
and stigma brownish.

FEMALE (ALLOTYPE). Measurements. HL 1.89; HW
2.05; SL 0.37; WL 4.8; TL 7.8.

Head. Broad, HW 1.08 x HL; scape short, SL 0. 19 x HL.
Eyes moderately convergent below, UFW 1.41 x LFW.
Clypeal length and apical width subequal; BCW 0.54 x CW;
BCW:COD:CAD:ASD:IAD = 13:12:6:5:7. OD:IOD:OOD =
5:12.5:9. Mandible broad, tridentate, inner tooth small.
Clypeus coarsely and closely punctate, frons a little more
coarsely punctate; interspaces dull, densely tessellate. Facial
fovea ending about *A of distance between eye and ocellus.

Thorax. As in male, but pleural punctures a little finer, no
sparser above than below.

Abdomen. Similar to that of male, with usual sexual dif-
ferences; no tumescence on sternum 3.

Pilosity. Much as in male, but a little denser in all areas.

Color. Black; lower sides of face, mandible, clypeus except
black basal third, under side of scape and flagellum, legs, all
ferruginous. Side of face with broad yellowish macula ending
abruptly above at level of antennal sockets. Upper side of
scape and flagellum brownish. Tegula translucent brownish.
Wings clear, veins and stigma light brown. First tergum with
reddish areas laterally and basally.

TYPE MATERIAL

Holotype male and allotype: Khami, ZIMBABWE, 1 1 Dec.
1932 (no name), from collection of the National Museum of
Zimbabwe, deposited in SAM: Paratype: 19, Pretoria, Trans-
vaal, SOUTH AFRICA, 9 Jan. 1980 (, SJ. van Tonder; PRET).

ETYMOLOGY

Latin, acarus (mite) plus the suffix -phorus (to bear), in ref-
erence to the presence of a mite-bearing chamber at the base
of the first tergum.

14 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

Figures 27-31. H. (Alfkenylaeus) acariphorus: 27-28, frontal view of head of female and male (scale line = 1 .00 mm); 29-31, male sterna 7
and 8, genitalic capsule (scale line = 0.50 mm). Figures by R.A. DeNicola.

DISCUSSION

This species is easily recognized by the diagnostic characters
given above. The Zimbabwe specimens possess hypophal
mites in the modified basal sulcus of the first tergum.

Two additional females, which may be this species, have
been seen. Both are from Mombasa, KENYA, collected 12

Dec. 1982 by T.L. and R.T. Griswold, and are in Mr. Gris-
wold’s collection. They differ from the allotype in having the
first tergum ferruginous rather than black. The punctures
laterad on the metanotum are separated by about one-half a
puncture diameter, the basal area of the propodeum is very
weakly rugulose and the entire propodeum is matt. In one

Contributions in Science, Number 361

Sneliing: Ethiopian Hylaeine Bees 15

specimen the punctures of the second tergum are separated
by up to a puncture diameter and the punctures, instead of
being deep and sharply defined, slope upward to the tergal
surface along their posterior portions.

Possibly these represent another species, but this is un-
certain in the absence of males, and because there are so few
specimens that I have no idea of the limits of infraspecific
variation. For the time being it seems best to tentatively
assign these two Kenyan specimens to H. acariphorus. One
of these females has hypophal mites in the sulcus of the first
tergum.

Hylaeus (Alfkenylaeus) infulatus, new species

DIAGNOSIS

Female only: Terga 1-5 with complete apical pubescent fas-
ciae; clypeus immaculate and coarsely, contiguously to sub-
contiguously punctate in middle. Male unknown.

DESCRIPTION

FEMALE (HOLOTYPE). Measurements. HW 1.84; FIL
1 .7 1 ; SL 0.45; WL 4.4; TL 7.0 mm. Paratype: HW 1 .87; HL
1.74; SL 0.44; WL 4.3; TL 6.3 mm.

Head. Broad, HW 1.08 x HL; scape moderately long, SL
0.26 x HL. Eyes moderately convergent below, UFW
1.45 x LFW. Clypeus about as long as broad, sides regularly
divergent to broadest point, BCW 0.58 x CW; BCW:COD:
CAD:ASD:IAD= 32:24:17:13:18. OD:IOD:OOD = 11:33:
20. Frontal shield narrow, sides nearly straight and strongly
convergent above. First flagellar segment transverse, slightly
shorter than pedicel and about as long as second segment.
Entire clypeus coarsely and contiguously to subcontiguously
punctate, interspaces tessellate and moderately shiny, bot-
toms of punctures shiny. Remainder of front of head similar,
but interspaces of frons and vertex shiny, becoming more
distinctly tessellate in preoccipital area. Gena moderately
shiny between contiguous moderate to coarse punctures.

Thorax. Pronotal collar sharply carinate across front, Ca-
rina extending across front of lateral lobe. Mesoscutum about
1.6 x broader than long. Scutellum flat, about 0.36 x length
of mesoscutum. Metanotum, in profile, weakly convex, its
dorsum on same level as scutellum, about one-half as long
as scutellum. Basal face of propodeum sharply curved into
declivitous face; basal triangle sharply marginate. Side of
pronotal collar shiny between irregularly spaced fine punc-
tures; mesoscutum moderately shiny between subcontiguous
coarse punctures; scutellum similar but a little shinier and
punctures distinctly more separated in middle; metanotum
coarsely and subcontiguously punctate, interspaces moder-
ately shiny. Mesopleuron moderately shiny between coarse,
subcontiguous punctures; metapleuron slightly shiny be-
tween subcontiguous to close moderate punctures. Propodeal
triangle dull, appearing almost granulose, and sharply retic-
ulorugose; stigmatal area and side dull, moderately rugoso-
punctate; disc dull, finely rugosopunctate.

Abdomen. Tergum 1, in dorsal view, about 1.4 x broader
than long; basal sulcus extending above middle of anterior

face; disc shiny between coarse, subcontiguous punctures;
pregradulus of second tergum moderately shiny between sub-
contiguous to close, fine to moderate punctures; disc of ter-
gum 2 moderately shiny between subcontiguous to close
moderate to coarse punctures. Remaining terga moderately
shiny and finely tessellate between close to sparse fine punc-
tures.

Pilosity. Lower frons with conspicuous long, subappressed,
plumose hairs near antennal sockets; pronotal collar with
dense pubescent fascia; terga 1-5 with complete apical pu-
bescent fasciae; tergum 1 without erect hairs across summit
of anterior face.

Color. Black; antenna (lighter beneath), tegulaand legs dark
brownish. The following pale yellowish: large lateral face
mark, filling area between clypeus and eye, ending at level
of lower margin of antennal socket; small tegular spot; basal
spot on protibia and larger spot on metatibia. Wings clear,
veins and stigma brownish.

TYPE MATERIAL

Holotype female: Konkoyo, 22 km W Kebemer, SENEGAL,
4 Aug. 1979 (A. Pauly, #10) in GEMB. Paratype female:
Dingasso, near Bobo, UPPER VOLTA, 28 Sept. 1979 (A.
Pauly), on Ziziphus mauritiana, in LACM.

ETYMOLOGY

From Latin, adorned with a fillet or band, referring to the
abdominal fasciae.

DISCUSSION

Only the two female specimens are known. The paratype is
very similar to the holotype: HW 1.07 x HL; UFW 1.43 x
LFW; BCW 0.59 x CW; BCW:COD:CAD:ASD:IAD = 33:
23:16:13:21. OD:IOD:OOD = 10:34:22.

Although very similar to H. namaquensis, females of H.
infulatus are more coarsely punctate; this is especially evident
on the clypeus, as noted in the key. The male of H. infulatus
is unknown but probably will run to H. namaquensis in the
key above.

Hylaeus ( Alfkenylaeus ) namaquensis Cockerell

Figures 32-36

Hylaeus namaquensis Cockerell, 1942:12-13. 9 8. SOUTH
WEST AFRICA: Aug., Jan. 1930 (R E. Turner) (BMNH)
[examined].

DISCUSSION

This species is easily recognized, in the female by the com-
bination of coarse punctation, extending to the second ter-
gum, black clypeus, but with two lateral marks on face and
pubescent fasciae on the second to fifth terga. The male has
the lower half of the face yellow, the third sternum without
a glabrous swelling, coarse punctation, and third to sixth terga
with preapical pubescent fasciae; the fasciae of the fourth and

16 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

Figures 32-36. H. ( Alfkenylaeus ) namaquensis: 32-33, frontal view of head of female and male (scale line = 1.00 mm); 34-36, male sterna
7 and 8 (ventral and lateral), genitalic capsule (scale line = 0.50 mm). Figures by R.A. DeNicola.

following segments may be absent due to abrasion. In this
species the propodeum is sharply and densely tessellate, with
fine, dense punctures; the basal triangle is rugosoreticulate at
the base.

SOUTH WEST AFRICA: 10<3<3, 1422, Aug., Jan. 1930 (RE.
Turner; BMNH, inch cotypes); 1 <3, Aug., Dec. 1929 (R E.
Turner; BMNH); IS, Windhoek, 12 Dec. 1933 (J. Ogilvie;
BMNH); 1<3, Kaoko Otavi, Mar. 1926 (no name; SAM). One

Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees 17

additional female, surely mislabelled, is in the BMNH: Mah-
datha, 60 mi. NE Mecca, ARABIA, Jan. 1945 (B.P. Uvarov).

Hylaeus ( Alfkenylaeus ) psaenythioides,
new species

Figures 37-40

DIAGNOSIS

Male only: Face with transverse yellow band between inner
orbits above clypeal base; punctation coarse. Female un-
known.

DESCRIPTION

MALE (HOLOTYPE). Measurements. HL 1 .68; HW 1 .74;
SL 0.68; WL 4.3; TL 6.5 mm.

Head. Broad, HW 1 .06 x HL; scape moderately long, twice
longer than wide, SL 0.40 x HL. Eyes strongly convergent
below, UFW 1.62 x LFW. Clypeus slightly longer than wide
at apex, sides evenly divergent from base; BCW 0.58 x CW;
BCW:COD:CAD:ASD:IAD = 1 1:8:7:5:5:5. OD:IOD:OOD =
5:11:8. Cephalic punctures uniformly coarse, about 0.08 mm
diam., often irregularly shaped, subcontiguous, interspaces
polished; genal punctures a little smaller, interspaces less
shiny, faintly tessellate.

Thorax. Pronotal collar with thin crest along anterior mar-
gin, reduced in middle, extended laterad along front of pro-
notal lobe nearly to lower margin of lobe; humeral ridge
sharp. Mesoscutum about 1.3 x wider than long. Scutellum
flat, about 0.4 x length of scutum. Metanotum half as long
as scutellum, anterior margin raised above posterior margin
of scutellum, sloping to propodeal base. Basal area of pro-
podeum oblique, evenly rounded onto posterior face; basal
triangle almost entirely on basal face, sharply marginate; me-
dian groove deep and narrow. Entire thorax with coarse,
subcontiguous punctures, those of mesoscutum about 0.06
mm diam., on mesopleura a little finer, propodeal punctures
coarser above than below; interspaces smooth and shiny and
dorsal areas, lightly tessellate and slightly shiny on pleura
and sides of propodeum.

Abdomen. Enlarged basal sulcus about % length of basal
face of tergum 1 ; apical impunctate band of tergum 1 broad,
sharply depressed; tergum 2 with apical impunctate band
broader, more depressed, especially at sides; tergum 3 with
apical impunctate band about as broad as on second, lightly
depressed; sternum 3 with low, inconspicuous shiny swelling
at base. First two terga coarsely punctate, punctures about
0.06 mm diam., interspaces smooth and shiny; remaining
terga slightly shiny, transversely lineolate and with sparse,
irregular, fine punctures; sterna shiny, very lightly tessellate
and with scattered fine punctures which are coarser than on
tergum 3. Sternum 7 with apical process expanded distally,
apical margin rounded; sternum 8 with apical lobes narrow,
apices reflexed, with a few setae along distal margin; gono-
coxite evenly narrowed, apices not broadened.

Pilosity. Specimen apparently rubbed. Short simple hairs

on front of head, a few longer, plumose hairs around antennal
sockets, upper inner orbits, occipital margin, and head. Tho-
racic dorsum with short simple erect hairs, longer, plumose
hairs at wing bases and sides of scutellum, metanotum, and
stigmatal area; pronotal collar with band of dense, appressed,
short plumose hairs and pronotal lobe margined by similar
hairs; pleura with only short simple hairs (longer, plumose
hairs may have once been present); propodeum with a few
long, plumose hairs at sides of posterior face. Tergum 1 with
moderately long simple hairs in sulcus, shorter simple hairs
at sides and on disc; tergum 2 with similar, longer hairs on
disc and sides; third and following terga with longer, more
abundant simple hairs. Sterna with sparse, long, simple hairs.
Tergum 1 with short, dense, appressed, plumose hairs on
each side of apical margin.

Color. Black, abdomen obscurely reddish basally and ven-
trally; mandible, labrum, apical fourth of clypeus, lower sides
of face, and most of legs reddish; underside of scape and
flagellum dull yellowish red, dorsal surfaces brownish. Meso-
and metafemora and tibiae mostly brownish. Tegula trans-
parent brownish. Wings clear, veins light brown, stigma dark-
er. Supraclypeal area and adjacent side of face light yellowish,
so that face has transverse yellow band (Fig. 37).

TYPE MATERIAL

Holotype male: 13 mi. S Malindi, KENYA, 26 May 1967
(C.D. Michener), in UKAN.

ETYMOLOGY

This name was suggested by the presence of the transverse
facial mark, as in the Neotropical bee genus Psaenythia (An-
drenidae), to the name of which is added the suffix, -oides,
resembling.

DISCUSSION

The species is easily recognized by the transverse facial mark,
apparently unique among the hylaeines of the Ethiopian Re-
gion. The female possibly will be similarly marked, though
it seems more likely that the supraclypeal area will be dark
in this sex.

Subgenus uncertain

Although Hylaeus arnoldi will key to the subgenus Alfken-
ylaeus it is not, in my opinion, a member of that subgenus.
In particular, I am impressed by the very different male
sternum 8 (Fig. 44) and the shape of the male gonocoxite
(Fig. 45). At present H. arnoldi does not fit within any of the
existing subgenera. I am, however, presently unwilling to
erect a monotypic subgenus for this species.

Hylaeus arnoldi (Friese)

Figures 41-45

Prosopis arnoldi Friese, 1913:574. <3. ZIMBABWE: Bula-
wayo, 28 Sept. 1912 ( G . Arnold) (MNHU) [examined].

18 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

Figures 37-40. Male, H. ( Alfkenylaeus ) psaenythioides: 37, frontal view of head (scale line = 1.00 mm); 38-40, sterna 7 and 8 (ventral and
lateral views), genitalic capsule (scale line = 0.50 mm). Figures by R.A. DeNicola.

Prosopis xanthopus Alfken, 1914:107. 2. ZIMBABWE: Bu-
lawayo, 28 Sept. 1912 {G. Arnold) (MNHU) [examined].

DISCUSSION

Bridwell (1919) correctly recognized that Alfken’s P. xan-
thopus was a synonym of P. arnoldi. His assignment of this

bee to Deranchylaeus appears to have been based on the
descriptions alone. The male terminalia are different from
those of Deranchylaeus; the lobules of sternum 7 lack teeth,
sternum 8 is profoundly bilobed and the gonocoxites are
sharply narrowed toward their apices and extend well beyond
the level of the apices of the penis valves. The shape of

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Snelling: Ethiopian Hylaeine Bees 19

sternum 8 will immediately separate this species from those
assigned here to the new subgenus Alfkenylaeus. The female
cannot be separated from those of the few species of Alfken-
ylaeus.

SPECIMENS EXAMINED

ZIMBABWE: 18, 19, Bulawayo, 28 Sept. 1912 (G. Arnold;
MNHU, types of P. arnoldi and P. xanthopus, respectively);
ASS, same data as P. arnoldi type (MNHU, SAM); 18, Bu-
lawayo, 21 Sept. 1913 ( G . Arnold; BMNH); 18, Bulawayo,
19 Oct. 1924 (R.H.R. Stevenson; AMNH); 18, Bulawayo, 5
Oct. 1924 (no name; SAM). SOUTH WEST AFRICA: 18,
Kaoko Otavi, Mar. 1926 (no name; SAM); 18, Okosongo-
mingo Farm No. 149, 59 km ESE Otjiwarongo, 17 Nov.
1972 (C.L. Hogue; LACM); 19, Karasburg, 850 m elev., 24
Sept. 1967 (Ross and Stephen; CAS).

Subgenus Mety/aeus Bri dwell

Mety/aeus Bridwell, 1919:131. Type-species: Metylaeus crib-
ratus Bridwell, 1919; original designation.

DIAGNOSIS

Preoccipital carina present; anterior and lateral faces of mes-
epistemum separated by a carina on lower half; metanotum
sharply marginate at sides, usually produced as spiniform
process.

DESCRIPTION

(1) Mandible broad, bidentate at apex. (2) Clypeus without
preapical depression, with dense to contiguous moderate
punctures. (3) Preoccipital carina present. (4) Pronotal collar
with anterior carina which extends laterad to posterior lobe.
( 5) Mesepistemum carinate between anterior and lateral faces
in lower half. (6) Lateral, oblique, and transverse propodeal
carinae present, strong; basal triangle coarsely areolate. (7)
Sulcus of tergum 1 broad, less than half as long as basal face.
(8) Tergum 2 with punctiform lateral fovea. (9) Male second
and third sterna simple. (10) Male sternum 7 with basal
lobule absent or poorly defined, without lateral teeth (Fig.
46). (11) Male gonocoxite broad, blunt, ending slightly be-
yond level of apex of penis valves, with numerous long,
barbed hairs (Fig. 50).

Labral tubercle present in both sexes, not well defined in
male; I AD about 1.2 x COD; subantennal sutures about
1.5 x ASD; upper end of female facial fovea ending near
inner eye margin; first flagellar segment broader than long,
about one-half as long as second segment; scutellar processes
present (Figs. 54-55); frontal shield unusually high and short;
terminating abruptly a little above level of antennal sockets;
dark, slender species with dense to close moderate punctures,
usually including first one or two terga.

DISCUSSION

Bridwell (1919) proposed Metylaeus as a genus, based in large
part on the conspicuous modifications of the scutellum and

metanotum. He had available for study both sexes of the
type species, but had seen no others. The only other species
of which he was aware were two species known from males
only: H. scutispinus (Alfken) and H. catalaucoides (Bridwell)
(H. catalaucoides was a new name for the improperly asso-
ciated male of H. bouyssoui, a species which Bridwell as-
sumed to belong to his subgenus Deranchylaeus of Hylaeus );
these he knew only from descriptions. Since all of these were
known to possess both scutellar and metanotal spines, the
presence of such spines was assumed to be characteristic of
the new genus. Samples of additional species have negated
the significance of these spines as a generic character.

In the females of H. bouyssoui and H. scutispinus the scu-
tellum is weakly depressed posteromesally, but otherwise is
simple. The median area of the metanotum is marked by a
sharp oblique carina on each side; the posterior ends of the
carinae are joined by a transverse carina, but there are no
spines. In females of these species the preoccipital carina is
weak, and these are superficially similar to some species of
Deranchylaeus. The male of H. scutispinus has well-devel-
oped spines on both scutellum and metanotum and is oth-
erwise similar to H. cribratus; the two must be placed in the
same group.

Thus, the distinctions between Metylaeus and Deranchy-
laeus are less clear-cut than once seemed to be the case. I
believe that they are to be treated as related subgenera of
Hylaeus. Popov (1939) suggested that the Philippine sub-
genus Hoploprosopis ought not be separated from Metylaeus.
As I have shown elsewhere (Snelling, 1969), the two are
readily separable and presumably not at all closely related.

Of the species listed below, H. spiniger (Benoist) is known
only from Madagascar and is not treated here.

SYNONYMIC LIST OF SPECIES

bouyssoui (Vachal)
cribratus (Bridwell)

=catalaucoides Bridwell, NEW SYNONYMY
=rugiceps Friese, NEW SYNONYMY
=semlikiensis Cockerell, NEW SYNONYMY
gaullei (Vachal)
scutispinus (Alfken)
spiniger (Benoist)

KEY TO SPECIES OF METYLAEUS

1. Antenna 1 2-segmented; female 2

- Antenna 13-segmented; male 5

2. Scutellum simple; metanotum with laterally marginate

median area, but no spines 3

- Scutellum and metanotum deeply excavated and with
posteriorly directed lateral spines . . cribratus (Bridwell)

3. Tergum 1 finely and closely punctate, second densely tes-
sellate and impunctate or nearly so; tergum 2 without
apical pubescent fascia; clypeus blackish 4

- First and second terga coarsely and subcontiguously to

densely punctate; tergum 2 with complete apical pubes-
cent fascia; clypeus partly ferruginous

scutispinus (Alfken)

20 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

Figures 41-45. H. { Deranchylaeus ?) arnoldi: 41-42, frontal view of head of female and male (scale line = 1.00 mm); 43-45, male sterna 7
and 8 (ventral and lateral views), genitalic capsule (scale line = 0.50 mm). Figures by R.A. DeNicola.

4. Interspaces of tergum 1 slightly shiny, obviously sculp-
tured; second tergum virtually impunctate; first two terga

without subdecumbent to suberect hairs

bouyssoui (Vachal)

- Interspaces of tergum 1 smooth, subpolished; second ter-

gum with sparse to scattered minute punctures; first two
terga with scattered subdecumbent to suberect hairs, es-
pecially laterad gaullei (Vachal)

5. Scape, flagellum and metabasitarsus black; punctures of
tergum 2 much finer than those of first (rarely, punctures

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Snelling: Ethiopian Hylaeine Bees 21

may be very obscure), apical margin not, or weakly, de-
pressed cribratus (Bridwell)

- Scape and flagellum reddish, metabasitarsus reddish to
yellow; tergum 2 nearly as coarsely and closely punctate

as first, apical margin sharply depressed

scutispinus (Alfken)

Hylaeus ( Metylaeus ) bouyssoui (Vachal)

Figure 53

Prosopis Bouyssoui Vachal, 1899:535. $ only. GABON:

N’Doro, 15 Sept, and 12 Nov. 1898 (J. Bouyssou) (MNHN)

[examined].

DIAGNOSIS

Female only: Scutellum and metanotum without sublateral
spiniform processes; tergum 1 tessellate and moderately shiny
between fine, close punctures, middle impunctate or nearly
so. Male unknown.

DESCRIPTION

FEMALE. Measurements. HL 1.56; HW 1.63; TL 5.6,
WL 4.3.

Flead. Slightly wider than long; HW 1.04 x HL; scape
short, SL 0.32 x HL, SL 2.80 x SW. Eyes moderately
convergent below, UFW 1.48 x LFW. Clypeus broad, CW
1.05 x CL; BCW 0.55 x CW, 2.20 x ASD, 1.57 x IAD,
1.37 x COD. Clypeal punctures shiny within, moderate,
contiguous to subcontiguous, round, shallow, interspaces
slightly shiny; supraclypeal area with contiguous, slightly
coarser punctures; paraocular areas similar to clypeus; sides
of supraclypeal shield moderately flared, margins effaced,
disc rugulose, apical width about 0.25 x ASD; punctures of
vertex and occiput a little coarser, more regular in shape,
shiny within; gena slightly shiny, with moderate to coarse
contiguous punctures, shiny within. Fovea ending slightly
nearer eye than ocellus.

Thorax. Carina of pronotal collar sharp, disc with dull
interspaces between moderate, subcontiguous punctures.
Mesoscutum about 1.15x wider than long. Scutellum flat,
about 0.34 x length of mesoscutum. Metanotum with me-
dian, laterally and posteriorly carinate trapezoidal area, in-
terior of which is irregularly, finely rugulose; lateral areas dull
and tessellate near trapezoid, finely and contiguously punc-
tate at extreme side. Mesoscutum dull between moderate,
contiguous to subcontiguous punctures; scutellum weakly de-
pressed in middle, more strongly so posteriorly, depressed
area with fine to moderate, subcontiguous punctures which
become fine and contiguous posteriorly, lateral areas with
irregularly spaced, mostly moderate punctures; mesopleuron
dull between coarse, contiguous to subcontiguous shallow
punctures which are shiny within; metapleuron dull, mod-
erately, contiguously punctate. Side of propodeum slightly
shiny, finely, closely and irregularly rugulose; stigmatal and
discal areas coarsely rugulose; basal triangle coarsely, quad-
rately areolate.

Abdomen. Tergum 1 about 1.5 x wider than long, disc
moderately shiny between fine, subcontiguous to dense punc-

tures, impunctate along midline; pregradulus of tergum 2
moderately shiny between scattered minute punctures, grad-
ulus weakly impressed, disc moderately shiny between very
obscure, sparse, minute punctures, margin not depressed in
middle.

Pilosity. Propodeum pollinose; first and second terga with-
out apicolateral pubescent fasciae.

Color. Black. A pair of submedian spots on pronotal collar,
part of posterior pronotal lobe and basal spot on protibia,
yellowish. Antenna and legs brownish, flagellum paler be-
neath. Wings slightly brownish, veins and stigma dark brown.

TYPE MATERIAL

Described from two females (15 Sept, and 12 Nov. 1898)
from N’Doro, GABON, collected by J. Bouyssou. The male
described by Vachal is not conspecific. Of the original two
females of P. bouyssoui, one is in the Paris Museum and
bears a red TYPE label and another label, in Vachal’s hand:
“Bouyssoui/Vach.” This specimen is here selected as the
lectotype.

DISCUSSION

Because the female does not possess scutellar and metanotal
spines, Bridwell (1919) placed P. bouyssoui in his subgenus
Deranchylaeus of Hylaeus; the male was recognized to belong
to Metylaeus. In the original description of the male, Vachal
stated that the metanotal spines were triangular. With this
distinction between Vachal’s male specimen and those which
he had described as M. cribratus, Bridwell renamed the Va-
chal male as M. catalaucoides.

The female of H. bouyssoui is very similar to that of H.
gaullei but is smaller, the mesoscutum is dull between con-
tiguous to subcontiguous punctures and the first tergum is
moderately shiny and distinctly tessellate between fine, close
punctures, except along the essentially impunctate midline.

MATERIAL EXAMINED
Only the lectotype has been seen.

Hylaeus ( Metylaeus ) cribratus (Bridwell)

Figures 46-47, 51-52

Metylaeus cribratus Bridwell, 1919:131-133. <5 9. NIGERIA:
Ibadan, Aug.-Sept. 1914 (J.C. Bridwell) (USNM) [ex-
amined].

Metylaeus catalaucoides Bridwell, 1919:133. <5. GABON:
N’Doro, 30 Sept. 1 898 ( J . Bouyssou) (MNHN) [examined].
New name for Prosopis Bouyssoui Vachal, <3, not 9. NEW
SYNONYMY.

Prosopis rugiceps Friese, 1921:1 105-1 106. 9. ZAIRE: Duma,
Ubangi District, 20 Oct. 1910 ( Schubotz ) (type depository
unknown). NEW SYNONYMY.

Metylaeus semlikiensis Cockerell, 1936:1 1. 9. ZAIRE: Sem-
liki Valley, 16 Aug. 1914 (J. Bequaert) (AMNH) [exam-
ined], NEW SYNONYMY.

DISCUSSION

The male which Vachal (1899) described as that of Prosopis
bouyssoui was correctly recognized by Bridwell (1919) as not

22 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

49

Figures 46-50. Males, H. ( Metylaeus ) spp.: 46-47, sterna 7 and 8, H. cribratus; 48-50, sterna 7 and 8, genitalic capsule H. scutispinus (scale
line = 0.25 mm).

being conspecific with the female. Accordingly, he renamed
the male as Metylaeus catalaucoides. The name is based on
that of the ant genus Cataulacus and is, therefore, misspelled.
The erroneous spelling evidently did not originate with i Ind-

well. for Vacha! wrote “fere sicut in Catalauco reticulata Sm.
. . . .” The point is moot, however, since this name is a junior
synonym of H. cribratus.

Vachal’s male is from N’Doro and is in the Paris Museum.

Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees 23

It bears a label in Vachal’s hand: “Bouyssoui/Vach.” Inas-
much as this is the type of M. catalaucoides, I have attached
to it a red label: “TYPE/ Metylaeus/ catalaucoides/BRlD-
WELL 1919.”

Although no type material of P. rugiceps has been exam-
ined, nothing in the description would indicate this to be
anything other than H. cribratus; Friese may not have been
aware of Bridwell’s species when he described P. rugiceps.
The specimens from Cameroon and Uganda were identified
as P. rugiceps by Alfken, who may have seen type material
of this name. Cockerell’s M. semlikiensis is identical to other
females from Zaire which form part of a continuous series
of variants to typical H. cribratus. Cockerell stated otherwise,
but there are remnants of a hair band along the margin of
the first tergum of the type of M. semlikiensis.

MATERIAL EXAMINED

NIGERIA: 93,3, 1522, Oloke Meji, Ibadan, Aug.-Sept. 1914
(J.C. Bridwell; USNM) (type series of M. cribratus ); 13, 12,
Lagos, 18 Aug. 1966 (C.D. Michener; UKAN). CAMER-
OON: 12, Akoafim, no date (S.G. Tessmann, No. 15-31;
SAM). GABON: 13, N’Doro, 30 Sept. 1898 (/. Bouyssou;
MNHN) (type of M. catalaucoides). ZAIRE: 12, Semliki Val-
ley, 16 Aug. 1914 (J. Bequaert; AMNH) (type of M. semli-
kiensis)-, 13, 61 mi. E Kenge, 5 Aug. 1957; 13, 422, 18 mi. W
Luanza, 1 300 m elev., 1 6 Jan. 1958; 13, 39 mi. NE Lusambo,
12 Aug. 1957; 13, 222, Irangi, 900 m elev., Luhoho R., 10
Sept. 1957; 12, 39 km SWalikale, 700 m elev., 25 Dec. 1957;
12, 33 mi. SW Kamituga, 675 m elev., 17 Aug. 1957 (all E.S.
Ross & R E. Leech; CAS). UGANDA: 422, no further data
(MNHU). ANGOLA: 13, near Kasai R., July 1931 (T.D.A.
Cockerell; BMNH).

Hylaeus ( Metylaeus ) gaullei (Vachal)

Prosopis Gaullei Vachal, 1899:536. 2. GABON: Mouny, no
further data (MNHN) [examined].

DIAGNOSIS

Female. Scutellum and metanotum without sublateral spines;
tergum 1 subpolished, with moderate, irregularly spaced
punctures. Male. Unknown.

DESCRIPTION

FEMALE. Measurements. HL 1.80; HW 1.87; WL 5.3;
TL 6.6 mm.

Head. Broad, HW 1.03 x HL; scape short, SL 0.26 x HL;
SL 2.54 x SW. Eyes moderately convergent below, UFS
1.34 x LFW. Clypeus as broad as long; BCW 0.63 x CW,
2.50 x ASD, 1.36 x IAD, 1.67 x COD. Clypeus weakly de-
pressed on each side of middle, dull between shallow, mod-
erate, subcontiguous to dense punctures which are shinier

than interspaces; supraclypeal area with coarse, contiguous
punctures; paraocular area similar to clypeus, but punctures
mostly subcontiguous and interspaces slightly shiny; supra-
clypeal shield depressed in middle, with a few obscure mod-
erate to coarse punctures; frons slightly shiny between mod-
erate to coarse, contiguous to subcontiguous deep punctures;
vertex and occiput coarsely rugosopunctate, moderately shiny;
gena moderately shiny and finely lineolate between moderate
to coarse, contiguous to subcontiguous punctures which are
shiny within. Fovea ending a little less than halfway between
eye and ocellus.

Thorax. Carina of pronotal collar weak at side, slightly
depressed in middle, disc moderately shiny between fine to
moderate, irregularly spaced punctures. Mesoscutum about
1 .3 x wider than long. Scutellum flat, about 0.38 x length of
mesoscutum. Metanotum weakly convex, about half as long
as scutellum. Mesoscutum slightly to moderately shiny be-
tween moderate, mostly dense punctures; scutellum mod-
erately shiny, with very irregularly spaced, fine to coarse
punctures; metanotum in middle with large, laterally and
posteriorly carinate, trapezoidal shiny and irregularly rough-
ened area, lateral areas dull, moderately and contiguously
punctate; mesopleuron slightly shiny between shallow, flat-
bottomed, moderate to coarse, subcontiguous to dense punc-
tures which are moderately shiny within; metapleuron mod-
erately shiny, coarsely rugosopunctate. Side of propodeum
appearing dull because of dense, hoary pubescence, finely
rugosopunctate; stigmatal area and disc moderately rugo-
sopunctate.

Metasoma. Tergum 1 about 2.1 x wider than long, disc
subpolished, nearly impunctate along middle, otherwise with
close to sparse, fine punctures; tergum 2 moderately shiny
between minute sparse to fine punctures; remaining terga
duller, with scattered, obscure, ultraminute punctures.

Pilosity. Pronotal collar with conspicuous, though narrow,
transverse fascia; terga 1 and 2 without apicolateral fascia.

Color. Black. Legs, underside of flagellum and tegula brown.
Pronotal lobe with posterior yellowish blotch. Wings slightly
brownish, veins and stigma medium brown.

TYPE MATERIAL

Described from a single female. The type is in the Paris
Museum; there is no type label, but the data are correct, the
specimen matches the description and bears a label in Va-
chal’s hand identifying it as P. Gaullei. I have no doubt this
is the type and have affixed to it a red label: “TYPE? Prosopis/
Gaullei/VACHAL 1899.” The type is the only specimen ex-
amined of this species.

DISCUSSION

The original description is inadequate and Bridwell (1919)
assumed H. gaullei to be a species of Deranchylaeus close to

Figures 51-55. H. ( Metylaeus ) spp., dorsal view of scutellum, metanotum, and propodeal base: 51-52, female and male, H. cribratus; 53,
female, H. bouyssoui; 54-55, female and male, H. scutispinus (scale line = 0.50 mm, 51-52, 54-55 to same scale).

24 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees 25

H. dregei. It is, however, a Metylaeus close to H. bouyssoui.
The larger size, more sparsely punctate mesoscutum and shiny,
sparsely punctate first tergum will readily separate H. gaullei
from H. bouyssoui. The lack of scutellar and metanotal spines
will differentiate H. gaullei from H. cribratus.

Hylaeus ( Metylaeus ) scutispinus (Alflcen)

Figures 48-50, 54-55

Prosopis scutispina Alfken, 1914:195. 3. ZIMBABWE: Bu-
lawayo, 28 Sept. 1912 ( G . Arnold ) (MNHU) [examined].

DISCUSSION

The female of H. scutispinus lacks spines on the scutellum
and metanotum, although they are present in the male. Ad-
ditional features of the female are the ferruginous mandible,
labrum, and portions of the clypeus, the sharply and sub-
contiguously to densely punctate second tergum and the pres-
ence of a complete apical pubescent fascia on the second
tergum. The male is distinguished by the reddish antenna
and by the coarsely and closely punctate second tergum, the
apical margin of which is sharply depressed.

MATERIAL EXAMINED

UPPER VOLTA: 1329, 733, Bobo-Dioulasso, 20 Feb. 1980
(A. Pauly; GEMB), on Guiera senegalensis; 12, Mare-aux-
Flippopotames, 3 Oct. 1979 (A. Pauly; GEMB); 13, Dingasso
(near Bobo), 28 Sept. 1979 (A. Pauly; GEMB), on Ziziphus
mauritiana; 13, Kougny, 12 Feb. 1980 (A. Pauly; GEMB).
KENYA: 12, Diani Beach, Aug. 1951 ( N.L.H . Krauss;
BMNF1); 12, Guengere, Pungoue Valley, Mozambique, no
date {no name; MNHN). BOTSWANA: 12, Maun, 930 m
elev., 6 Nov. 1 976 {Ross and Stephen; CAS). SOUTH WEST
AFRICA: 13, 222, Kaoko Otavi, Mar. 1926 {no name; SAM).
ZIMBABWE: 13, Bulawayo, 28 Sept. 1912 (G. Arnold;
MNHU; type of P. scutispina ); 13, Bulawayo, 16 Feb. 1913
( R.H.R . Stevenson; SAM).

Nothylaeus Bridwell

Nothylaeus Bridwell, 1919:125-126. Type-species: Prosopis
heraldica F. Smith, 1853; original designation.

Nothylaeus , sub. Anylaeus Bridwell, 1919:129-130. Type-
species: Nothylaeus {Anylaeus) aberrans Bridwell, 1919;
original designation. NEW SYNONYMY.

DIAGNOSIS

Separable from all known hylaeine genera by the elongate,
slender mandible, with acuminate apex and without preapi-
cal tooth or with greatly reduced tooth.

DESCRIPTION

(1) Mandible elongate, apex acuminate, preapical tooth ab-
sent or greatly reduced. (2) Labral tubercle very broad, cov-
ering most of labrum, without median depression. (3) Ten-
torial pit at or (usually) below midpoint of clypeal length. (4)
First flagellar segment of male shorter than, or no longer

than, second. (5) Frontal shield present. (6) Lateral and oblique
propodeal carinae present, usually entire; propodeal groove
broad, shallow. (7) Sulcus of tergum 1 broad, less than one-
half as long as basal face. (8) Gradulus of tergum 2 gently
arched, slightly deflected laterad; lateral fovea broadly oval.

(9) Tergum 3 of male without sublateral pubescent fovea.

(10) Male sternum 7 bilobate, each lobe with proximal and
distal sublobes, lateral margin serrate or ciliate. (11) Male
sternum 8 with apical process short, broad, preapically ex-
panded; dorsal tubercle subapical or apical. (12) Male gono-
coxite narrow and elongate, extending much beyond apex of
aedeagus.

DISCUSSION

The precise status of Nothylaeus is somewhat dubious. Brid-
well (1919) proposed Nothylaeus as a genus and it has been
generally recognized as such, although Cockerell ( 1 936, 1 942)
seemed to be ambivalent. In originally characterizing Not-
hylaeus, Bridwell stressed the elongate, sharply pointed man-
dibles of both sexes and the greatly elongated gonocoxal apex
of the male.

The elongate gonocoxal apex is not unique to species of
Nothylaeus. This is a feature that occurs sporadically in some
species of African Hylaeus and in Hylaeus from other parts
of the world, as well. These are clear cases of morphological
character convergence.

The mandibular structure of both sexes is unique among
hylaeine bees. In very nearly all hylaeines, the mandibles of
both sexes are short and broad, the outer surface is marked
by distinct longitudinal ridges and grooves and the apical
margin is truncate to oblique, with one or more preapical
teeth. The mandible in Nothylaeus is remarkably elongate,
at least three times longer than broad at its midlength, there
are no distal and dorsal faces, as such, the preapical tooth is
absent or greatly reduced and the outer face of the mandible
lacks obvious grooves and ridges.

Nothylaeus, as a genus apart from Hylaeus, would be more
secure if additional supportive features could be found. The
robust habitus of Nothylaeus species is characteristic, but too
elusive to describe adequately. The head is relatively short
and broad and many parts of the body are extensively fer-
ruginous, but these features occur widely in Hylaeus. How-
ever, I am compelled to consider Nothylaeus separate from
Hylaeus, since the mandibular form is so consistent and
unique.

No such uncertainty prevails in the case of Anylaeus, pro-
posed as a subgenus of Nothylaeus by Bridwell (1919). Species
assigned to Anylaeus differed from those of Nothylaeus (s.s.)
in having the scutellum and metanotum modified in a man-
ner similar to those of the subgenus Metylaeus of Hylaeus.
This was true for both sexes of the species known to Bridwell
at that time. Now, species are known in which the modified
thoracic segments occur in the male but not the female. I
have here treated Anylaeus as a synonym of Nothylaeus. The
species of Nothylaeus will be revised in the second part of
this study.

26 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

LIST OF INCLUDED SPECIES NAMES

aberrans Bridwell, 1919
abyssinica (Alfken, 1905)
ameliae (Cockerell, 1942)
bevisi (Cockerell, 1917)
binotata (Alfken, 1914)
braunsi (Alfken, 1905)
dentiferella (Strand, 1912)
fortis Cockerell, 1936a
fumata (Strand, 1912)
gigas (Friese, 1911)
haemorrhoa Benoist, 1946*
heraldica ( F. Smith, 1853)
isochronous (Cockerell, 1936a)
junodi (Friese, 1911)
libericus Cockerell, 1936a
maculipes Cockerell, 1936a
magretti (Vachal, 1892)
montacuti Cockerell, 1942
neavei (Cockerell, 1942)
nigricans (Friese, 1913)
nyassana (Strand, 1912)
peringueyi Bridwell, 1919
rhodesicus Cockerell, 1942
rubrifacialis (Strand, 1912)
rubriplagiata (Cameron, 1905)
rufipedoides (Strand, 1911)
rufipicta (Strand, 1912)
sansibaribia (Strand, 1912)
simpsoni (Cockerell, 1 942)
subfortis Cockerell, 1 942
uelleburgensis (Strand, 1912)
ugandicus Cockerell, 1939
umtalicus Cockerell, 1936a
yoruba Bridwell, 1919

Calloprosopis, new genus

Type-species: Hylaeus magnificus Cockerell, 1942.

DIAGNOSIS

Body metallic blue in both sexes, female immaculate, male
with maculate clypeus; sulcus of first tergum narrow, ex-
tending nearly full length of basal face; female with elongate,
raised glabrous area at base of metatibia; male with gonobase
reduced, not forming cup at base of genital capsule.

DESCRIPTION

(1) Mandible stout, bidentate, apical margin oblique in fe-
male, transverse in male. (2) Labral tubercle prominent in
both sexes. (3) Tentorial pit at about midlength of clypeus.
(4) Pedicel, first and second flagellar segments about equal

* Known only from Madagascar.

in length, longer than broad. (5) Frontal shield present. (6)
Oblique propodeal carina absent, lateral carina very weak,
obvious only near its terminus; basal triangle almost entirely
on dorsal face; posterior groove deep and narrow. (7) Sulcus
of tergum 1 narrow, deep, extending almost entire length of
basal face. (8) Gradulus of tergum 2 broadly convex, pre-
gradulus much longer in middle than at sides; lateral fovea
absent, but spiracle in shallow depression. (9) Sternum 3 of
male with median, transverse, low swelling. (10) Male ster-
num 7 transverse, lobes reduced (Fig. 58). (11) Male sternum
8 with elongate distal process, apex transverse, dorsal tu-
bercle absent (Fig. 59). (12) Male genitalia massive, gono-
coxite robust, ending at about level of apex of penis valve
(Fig. 61).

Integument metallic blue, female without pale marks, male
with pale clypeal mark only; scape slender; female metatibia
with elongate, glabrous, basal ridge on outer side; male gono-
base forming a ring-like flange at base of genital capsule; male
volsella elongate, with prominent lateral tubercles.

ETYMOLOGY

The Greek kallos (beauty) plus Prosopis, an old generic name
for Hylaeus.

DISCUSSION

This genus closely resembles Hylaeus, but differs immedi-
ately from all known species of the Ethiopian and South
African regions by the metallic blue color. This character,
however, does appear in some Hylaeus groups in Australia
and the Philippine Islands. The presence of what appears to
be the basitibial plate in the female and the modifications of
the male terminalia are sufficient in my opinion to justify
recognition of Calloprosopis at generic level. The modifica-
tions of the male genital capsule, in particular, are unique
among the Hylaeinae. Among all Hylaeinae which I have
studied directly, and among those described and illustrated
by other workers, the gonobase is large and forms a cup-like
base to the genital capsule. In Calloprosopis the gonobase,
dorsally, projects into an emargination between the gono-
coxites; from this area it extends ventrad to form a heavily
sclerotized ring. The gonocoxites are heavily sclerotized and
are dorsoventrally broadened; they do not extend beyond the
apices of the penis valves. The volsellae are heavily sclero-
tized and the median lobes are elongate, with scattered small
tubercles on the outer faces. There is a deep longitudinal
groove along the entire length of the penis valves, the apices
of which are rather blunt and not as strongly deflected down-
ward as in Hylaeus. Sternum 8 of the male is typically hy-
laeine in appearance but lacks the notch at the base of the
apical process which is usually present in Hylaeus. Sternum
7, too, is typically hylaeine, but it is much broader than long;
the apodemes form a regular arc and the apical process is
quite short, with small lobes.

Most of the genitalic features are nothing more than ex-
treme modifications of conditions already present in other

Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees 27

hylaeines. The structure and orientation of the gonobase are
unique, however, and it is largely on this basis that Callo-
prosopis is given generic rank here. A cursory study of various
colletid genera in other subfamilies suggests that this pecu-
liarity may be unique within the family.

Calloprosopis magnified (Cockerell),
new combination

Figures 56-61

Hylaeus magnificus Cockerell, 1942:9-10. <3. KENYA: east
foot and slopes, Aberdare Mts., 7000-8500 ft. elev., 24-
27 Feb. 1911 (S’. A Neave) (BMNH) [examined].

Both sexes are immediately separable from all other known
hylaeines in the Ethiopian Region by their metallic color.
This species appears to be restricted to high elevations in
Kenya.

MATERIAL EXAMINED

KENYA. 1(3, east foot and slopes, Aberdare Mts., 7000-8500
ft. elev., 24-27 Feb. 1911 (S.A. Neave; BMNH, cotype); 1<5,
IS, Mt. Kinganop, 9000 ft. elev., Aberdare Range, cedar
forest, 27 Oct. 1934 (F. W. Edwards; BMNH); \S , Kerita,
2640 m elev., 38 mi. NW Nairobi, 16 Oct. 1957 ( E.S . Ross
& R E. Leech; CAS).

Psilylaeus, new genus

Type-species: Psilylaeus sagiops, new species.

DIAGNOSIS

Frontal shield absent; integument uniformly tessellate, with-
out obvious punctures; propodeum without defined basal
triangle; lateral fovea of second tergum broadly oval.

DESCRIPTION

( 1 ) Mandible short, sharply bidentate in male, weakly so in
female. (2) Labrum short and broad, male without tubercle,
female with elevated median tubercle which is weakly di-
vided in middle. (3) Tentorial pit slightly below midlength
of clypeus. (4) Male first flagellar segment transverse, shorter
than pedicel, as long as second, each shorter than third. (5)
Frontal shield absent. (6) Lateral and oblique propodeal ca-
rinae absent. (7) Sulcus of tergum 1 about one-half as long
as basal face. (8) Gradulus of tergum 2 weakly bowed, con-
cealed. (9) Tergum 3 with round sublateral pubescent fovea,
usually hidden under margin of second segment. (10) Male
sternum 7 bilobate, basal lobule setose, distal lobule with
apically hooked hairs (Fig. 64). (11) Male sternum 8 with
apical process broad, setose at margin; dorsal tubercle slightly
beyond midlength of apical process (Fig. 65). ( 1 2) Male gono-

coxite stout, blunt, not reaching level of apex of aedeagus
(Fig. 66).

Integument densely tessellate and dull, with sparse to scat-
tered, inconspicuous punctures on thoracic dorsum; front of
head dull between fine, contiguous punctures; eyes broadest
below midlength; IAD less than COD; frontal shield absent;
clypeal margins abruptly divergent in lower one-third; pos-
terior margin of pronotum much below dorsum of mesoscu-
tum, collar virtually absent except at sides; propodeum with
long, subhorizontal basal face.

ETYMOLOGY

The generic name combines the Greek psilos (bare or smooth)
with Hylaeus and refers to the virtually impunctate thoracic
dorsum, and especially to the smooth propodeum.

DISCUSSION

This genus is known to include only the type species, and is
known only from coastal South Africa in the vicinity of Cape
Town. The peculiarly smooth integument and unusual pro-
podeal structure are especially characteristic of this small bee.
In particular, the sutures which normally demark the pro-
podeal triangle are very weak and largely effaced.

Psilylaeus has been compared with various groups in hy-
laeines from Australia. In the key by Michener (1965), Psi-
lylaeus fails at the last couplet, since it does not agree with
either alternative ( Hylaeorhiza and Hylaeus). From Hylaeo-
rhiza, Psilylaeus differs in the bilobed, rather than acute,
glossa of the male and the outer apical angle of the hind tibia
is not obtuse. From Hylaeus, Psilylaeus differs (in the key)
in the structure of the propodeum. In the more recent key
to Australian genera by Houston (1975), Psilylaeus will run
to Hylaeus.

The depressed pronotum, without a well-defined collar,
occurs in two Australian subgenera of Hylaeus: Macrohy-
laeus and Hylaeteron. The former includes large, metallic
species with a long second submarginal cell in the forewing.
The known species of Hylaeteron are small, robust bees with
tridentate female mandibles, sharply reticulate propodeum
and exceptionally short subantennal sutures. Both of these
subgenera are known only from the Australian area.

Psilylaeus sagiops, new species

Figures 62-66

DIAGNOSIS

Same as generic diagnosis.

DESCRIPTION

MALE (HOLOTYPE). Measurements. HL 1 . 1 2; HW 1 .29;
SL 0.30; WL 3.50; TL 4.97 mm.

Head. Broad, HW 1.1 x HL; scape short, twice longer than

Figures 56-61. Calloprosopis magnifica: 56-57, frontal view of head, female and male (scale line = 1.00 mm); 58-61, male sterna 7 and 8,
genital capsule (lateral), genitalic capsule (dorsal and ventral) (scale line = 0.50 mm). Figures 61, 62 by R.A. DeNicola.

28 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees 29

wide, SL 0.27 x HL. Eyes strongly convergent below, UFW
1.55 x LFW, conspicuously broader below than above in
frontal view; in profile, broadest below midpoint, maximum
width 0.45 x EL. Clypeus slightly wider at apex than long,
epistomal sutures abruptly divergent in lower third; BCW
0.4 1 x CW x 1 . Clypeus and lower half of face slightly shiny,
densely tessellate, with scattered obscure shallow punctures;
immaculate areas of head duller, densely and finely punctate
and tessellate; gena slightly shiny, finely lineolate, with sparse,
shallow, fine punctures.

Thorax. Moderately robust, about 1 .4 x longer than wide.
Mesoscutum a little wider than long; in profile, anterior por-
tion convex, rising well above pronotum. Scutellum flat, on
same plane as posterior portion of scutum; median length
about 0.4 x that of scutum. Postscutellum flattened in pro-
file, sloping away from scutellum, median length less than
half that of scutellum. From above, sides of propodeum
strongly convergent distad, basal width almost twice apical;
in profile, basal face continuous with slope of postscutellum,
broadly rounded into, and longer than, posterior face; with-
out carinae or ridges. Slightly shiny, densely tessellate and
impunctate; mesopleuron and basal face of propodeum ob-
scurely lineolate; mesopleuron with sparse, fine punctures.

Abdomen. Widest beyond middle; apical width of tergum
1 greater than median length; sternum 3 with a pair of low,
shining prominences obliquely directed distad, on either side
of midline, highest at about middle; sternum 4 with a pair
of broad flattened, shiny callosities; sternum 5 with a similar,
but much smaller, pair; apex of sternum 6 broadly rounded.
Moderately shiny, finely transversely lineolate; all terga with
apical, nonsculptured band; tergum 1 with very fine scattered
punctures; tergum 2 more closely punctate, punctures larger;
tergum 3 similar to 2.

Terminalia. As described for the genus.

Pilosity. Very sparse; lower half of face with hairs short,
stiff; upper half with hairs much longer; scape with a few
moderately long hairs; underside of head with scattered long
hairs, especially in hypostomal area. Mesoscutum with very
short hairs only; scutellum and postscutellum with a few very
long hairs at sides; pleura and sides of propodeum with scat-
tered long hairs; propodeum otherwise with sparse, short
subappressed hairs. Terga with sparse, short hairs, appressed
on discs, longer and partially to fully erect laterad; progres-
sively longer on succeeding segments. Sterna with sparse erect
hairs, longest laterad.

Color. Blackish. Mandibles, except ferruginous apices; la-
brum; clypeus; transverse supraclypeal mark; face between
clypeus and eye, extending about halfway upward along inner
orbit; narrow line on underside of scape; narrow stripe on
sides of pronotal collar; pronotal lobe; apical spot on pro-
and mesofemora; basal mark on meso- and metatibia; all
light yellowish. Protibia and tarsi, mesotarsi, light ferrugi-
nous, apical tarsal segments brownish. Metabasitarsus whit-
ish on basal third, metatarsus otherwise brownish. Tibial
spurs whitish. Tegulae brownish. Wings uniformly light
brownish, veins and stigma darker.

FEMALE (ALLOTYPE). Measurements. HL 1.05; HW
1 . 1 1 ; SL 0.25; WL 3.30; TL 4.35 mm.

Head. Broad, HW 1.05 x HL. Eyes strongly convergent
below, UFW 1.52 x LFW, conspicuously broader below than
above; in side view, broadest part below midpoint, and
0.52 x EL. Mandibular apex braod, truncate, obscurely bi-
dentate. Clypeal shape as in male; BCW 0.46 x CW; BCW:
COD:CAD:ASD:IAD = 13:10:9:8:10. Scape and flagellum
as in male. Frontal line deep, terminating broadly at level of
lower margin of antennal sockets. Facial fovea terminating
at top of eyes, adjacent to eye margin. Surface sculpture as
in male.

Thorax. As in male.

Abdomen. As in male but lacking ventral modifications;
apical bands of terga broader.

Pilosity. As described for male but conspicuously shorter.
Color. Blackish. Minute basal spot on mandible; broad
longitudinal median stripe on clypeus; narrow stripe along
inner orbit, from lower end of eye to lower end of fovea;
narrow stripe on pronotal collar on each side, broadly inter-
rupted in middle; pronotal lobe; outer stripe on protibia;
basal spot on meso- and metatibia, all pale yellowish. Meta-
tibial spur, tegula, and wings as in male.

TYPE MATERIAL

All from Cape Province, SOUTH AFRICA: Holotype male,
Mossel Bay, 11 Oct. 1938 ( R.E . Turner). Allotype, same
locality, 12 Oct. 1938 (R.E. Turner). Holotype and allotype
in BMNH. Paratypes: 433, same data as holotype; 233, 22$,
same data as allotype; 13, same locality, 1 Oct. 1938 (R.E.
Turner)-, 12, same locality, Apr. 1933 (R.E. Turner)-, 733,
same locality, Mar.-Apr. 1930 (R.E. Turner ); 13, same lo-
cality, 7 Oct. 1941 (R.E. Turner)-, 12, same locality, 5 Dec.
1941 (R.E. Turner)-, 6 33, same locality, Nov. 1939 (R.E.
Turner ); 13, same locality, 12 Jan. 1940 (R.E. Turner)-, 2 33,
12, same locality, Jan. 1940 (no name)-, 12, Cape Town, 1
Apr. 1948 (no name); 12, Port Elizabeth, no date (N.L.H.
Krauss); 12, Worcester, Jan. 1934 (R.E. Turner). Paratypes
in BMNH, CORN, LACM, SAM, UKAN, USNM.

ETYMOLOGY

The specific epithet combines the Greek sagios (bag) with
ops (eyes), in allusion to the shape of the lower portion of
the eyes.

DISCUSSION

Variation is negligible in the limited amount of material
studied. The holotype is the largest specimen. The smallest
male has a head width of 0.98 mm and a wing length of 2.65
mm, with according allometric variations. Among the males
reduction in development of the glabrous processes of the
third to fifth sterna is relative to the size of the specimen.
Reduction begins with loss of the polished areas on the fifth
sternum and some diminution of those of the third and fourth.
Still smaller specimens lose next the pair on the fourth ster-
num, and the smallest male lacks modified areas on all three
sterna.

The few females examined are much more uniform. Head

30 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

Figures 62-66. Psilylaeus sagiops: 62-63, frontal view of head, female and male (scale line = 0.50 mm); 64-66, male sterna 7 and 8, gemtalic
capsule (scale line = 0.25 mm). Figures by R.A. DeNicola.

Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees 31

width varies from 1.05 to 1.13 mm and wing length from
2.59 to 3.32 mm. The smallest specimen has the head width
and head length equal. Smaller specimens tend toward loss
of punctation on the mesopleuron and first three terga, but
expression of this character is not uniformly correlated with
size.

The densely tessellate, impunctate integument, small size,
and lack of a defined supraclypeal area will readily separate
this species from all other Hylaeinae known from South Af-
rica. The trilineate face marks of the female are reminiscent
of the genus Allodape in the Anthophoridae.

ACKNOWLEDGMENTS

This project began in 1968 and has suffered many delays. I
must express my deep gratitude to all the curators who have
been so patient during the long time I have had material
under their care. For the loan of material from their respec-
tive institutions, I wish to thank: P.H. Amaud, Jr. (CAS),
G.C. Eickwort (CORN), G.R. Else and EH.H. Yarrow
(BMNH), H.E. Evans (MCZ), M. Favreau and J.G. Rozen,
Jr. (AMNH), A.J. Hesse (SAM), the late P.D. Hurd, Jr. and
G.I. Stage (USNM), S. Kelner-Pillault (MNHN), C.D. Mich-
ener (UKAN), J. Oelkhe (DEI), A. Pauly (GEMB), and E.
Pinhey (NMR). A very special gratitude is expressed to the
late Embrik Konigsmann (MNHU) for locating and making
available the critical types of the species described by Alfken,
Friese, and Strand.

My thanks, too, to my colleagues at the LACM; J.P. Don-
ahue, C.L. Hogue, and F.S. Truxal, for their advice and com-
ments over the years, both solicited and gratuitous. Some of
the illustrations were prepared by R.A. DeNicola, to whom
I remain grateful, as always.

LITERATURE CITED

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ten. Zeitschrift fur Hymenopterologie und Dipterologie
3:46-150.

. 1914. Zur Kenntnis der afrikanischen Prosopis-

Arten. Deutsche Entomologische Zeitschrift 1914:18 3—
197.

Benoist, R. 1 946 ( 1 945). Nouvelles especes d’Apides (Hym.)
de Madagascar. Societe Entornologique France, Bulletin
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. 1959. Les Prosopis de France. Cahiers des Natu-

ralistes, n.s. 15:75-87.

Bingham, C.T. 1903. On the Hymenoptera collected by
Mr. W.L. Distant in the Transvaal, South Africa, with
descriptions of supposed new species. Annals and Mag-
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. 1912. South African and Australian aculeate Hy-
menoptera in the Oxford Museum. Entomological So-
ciety of London, Transactions 1912:375-383.

Bridwell, J.C. 1919. Miscellaneous notes on Hymenoptera
with descriptions of new genera and species. Hawaiian
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Cameron, P. 1905. On some genera and species of Hy-

menoptera from Cape Colony and Transvaal. South
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. 1906. Descriptions of some new species of Hy-
menoptera from Pearston, Cape Colony. South African
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Cockerell, T.D. A. 1917. New records of Natal bees (second
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. 1919. Natal bees. Durban Museum, Annals 2: 1 89-

196.

. 1920. On South African bees, chiefly collected in

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. 1932. Descriptions and records of bees. CXXXV.

Annals and Magazine of Natural History, ser. 10, 10:
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. 1934. Some new or little known South African bees

of the genus Allodape in the British Museum. Annals
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. 1936a. African hylaeine bees. American Museum

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. 1936b. Descriptions and records of bees. CLVIII.

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631-638.

. 1939. Descriptions and records of bees. CLXXI.

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. 1942. Bees of the family Hylaeidae from the Ethi-
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Dathe, H.H. 1980. Die Arten der Gattung Hylaeus F. in
Europa (Hymenoptera: Apoidea, Colletidae). Zoolo-
gisches Museum Berlin, Mitteilungen 56:207-294.

Fabricius, J.C. 1793. Entomo/ogia systematica emendata
et aucta. Copenhagen, viii + 519 pp.

Friese, H. 1909. Die Bienen Afrikas. Zoologische und An-
thropologische Ergebnisse einer Forschungsreise im
Westlichen und Zentralen Sudafrika, Bd. 2, Lief. 1, X
Insecta, ser. 3, Jena, pp. 81-475.

. 1911. Die Maskenbienen der aethiopischen Region

(Prosopis, Hym.). Archiv fur Naturgeschichte 77:120-
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. 1913. Neue Bienenarten aus Afrika. Deutsche En-
tomologische Zeitschrift 1913:573-580.

. 1921. Ergebnisse der zweiten deutschen Zentral-

Afrika-Expedition 1910-1911, Apidae. Hamburgische
Wissenschaft Zeitung, pp. 1021-1 1 12.

Houston, T.F. 1975. A revision of the Australian hylaeine
bees (Hymenoptera: Colletidae). I. Introductory mate-
rial and the genera Heterapoides Sandhouse, Gephro-
hylaeus Michener, Hyleoides Smith, Pharohylaeus
Michener, Hemirhiza Michener, Amphylaeus Michener
and Meroglossa Smith. Australian Journal of Zoology >,
Suppl. Ser., 36:1-135.

. 1981. A revision of the Australian hylaeine bees

(Hymenoptera: Colletidae). II. Australian Journal of
Zoology, Suppl. Ser., 80:1-128.

Latreille, P.A. 1810. Considerations generates . . . des In-
sectes. Paris, 444 pp.

32 Contributions in Science, Number 361

Snelling: Ethiopian Hylaeine Bees

Linnaeus, C. 1758. Systema naturae. Editio decima, re-
formata. Stockholm, 824 pp.

Meade-Waldo, G. 1923. Hymenoptera, Fam. Apidae,
Subfam. Prosopidinae. In: P. Wytsman, Genera insec-
torum, 181:1-45.

Michener, C.C. 1965. A classification of the bees of the
Australian and South Pacific Regions. American Mu-
seum of Natural History, Bulletin 130:1-362.

. 1975. A taxonomic study of the African allodapine

bees (Hymenoptera, Anthophoridae, Ceratinini). Amer-
ican Museum of Natural History, Bulletin 155:67-240.

Popov, V.B. 1939. Subgeneric groupings of genus Prosopis
F. (Hymenoptera). Comptes Rendus (Doklady) de
I’Academie des Sciences de PURSS 25: 167-1 70.

Smith, F. 1 853. Catalogue of hymenopterous insects in the
collection of the British Museum. /. London: British Mu-
seum, 197 pp.

Sneliing, R.R. 1966a. Studies on North American bees of
the genus Hylaeus. 1. Distribution of the western species
of the subgenus Prosopis with descriptions of new forms
(Hymenoptera: Colletidae). Contributions in Science, no.
98, 18 pp. Natural History Museum of Los Angeles
County.

. 1966b. Studies on North American bees of the

genus Hylaeus. 2. Description of a new subgenus and
species (Hymenoptera: Colletidae). Washington Biolog-
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. 1966c. Studies on North American bees of the

genus Hylaeus. 3. The Nearctic subgenera (Hymenop-
tera: Colletidae). Southern California Academy of Sci-
ences, Bulletin 65:164-175.

. 1 968. Studies on North American bees of the genus

Hylaeus. 4. The subgenera Cephalylaeus, Metziella and
Hylaeana (Hymenoptera: Colletidae). Contributions in
Science, no. 144, 6 pp. Natural History Museum of Los
Angeles County.

. 1969. The Philippine subgenus Hoploprosopis of

Hylaeus (Hymenoptera: Colletidae). Contributions in
Science, no. 171,5 pp. Natural History Museum of Los
Angeles County.

. 1 970. Studies on North American bees of the genus

Hylaeus. 5. The subgenera Hylaeus, s. sir., and Para-
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Science, no. 180, 59 pp. Natural History Museum of Los
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5:745-768.

Accepted 14 November 1984.

Contributions in Science, Number 361

Sneliing: Ethiopian Hylaeine Bees 33

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Number 362
13 June 1985'

CONTRI

frafiNttraftY* A •• • . ■)

at|N T-jr :»/•' .• I ,• ' » •> ! i.,. ' , . ;

REVIEW OF THE LATE PLEISTOCENE AVIFAUNA
FROM LAKE MANIX, CENTRAL MOJAVE
DESERT, CALIFORNIA

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Craig C. Black, Museum Director

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Errol Stevens
Gary D. Wallace
Edward C. Wilson

REVIEW OF THE LATE PLEISTOCENE AVIFAUNA
FROM LAKE MANIX, CENTRAL MOJAVE
DESERT, CALIFORNIA

George T. Jefferson1

ABSTRACT. Late Pleistocene lacustrine clay, silt, and nearshore
sand deposits of Lake Manix, Mojave Desert, California, have yield-
ed 1 39 specimens of fossil birds, including several articulated partial
skeletons. Six extinct and 19 extant taxa are represented. The avi-
fauna includes the first recognized fossil gulls, Larus oregonus and
Larus sp., from Pleistocene terrestrial deposits in the Southwest.
Gavia cf. G. arctica (Arctic Loon), Phalacrocorax macropus (extinct
cormorant), Mergus cf. M. merganser (Common Merganser), the
extinct chronoclinal subspecies Fulica americana shufeldti (Shu-
feldt’s American Coot), and cf. Actitis (sandpiper) are new to the
Pleistocene record of the region. The avifauna is part of the Ran-
cholabrean Camp Cady Local Fauna, and has most taxa in common
with the assemblage from Fossil Lake, Oregon. Habitat preferences
of comparative extant taxa and inferred depositional environments
indicate extensive reedy marshlands, beach flats, and open water
lacustrine conditions.

INTRODUCTION

Since H. Howard’s description of the Lake Manix avifauna
(Howard, 1955), many additional specimens of fossil birds,
including articulated partial skeletons, have doubled the
number of taxa known from the site. These additional ma-
terials add significantly to an understanding of the late Pleis-
tocene avifauna of the inland Southwest. The first fossil gulls,
Larus oregonus and Larus sp. and other extant and extinct
taxa including Gavia cf. G. arctica, Phalacrocorax macropus
(extinct), Mergus cf. M. merganser, Fulica americana cf.
shufeldti (extinct), and cf. Actitis are new to the Pleistocene
record of the region. The Lake Manix avifauna is taxonom-
ically most similar, in the abundance of inland lacustrine
forms, to the Wisconsinan age assemblage from Fossil Lake,
Oregon. Other comparable late Pleistocene assemblages are
known from China Lake, McKittrick, Rancho La Brea, and
Carpintaria, California (Table 1). New radiometric dates (Bi-
schoff pers. comm., 1982) indicate that most of the material
from Manix is older than 200 Kyr (thousand years) BP (be-
fore present).

J.P. Buwalda of the University of California, Berkeley
(UCB), recovered the first bird fossils from Lake Manix dur-

Contributions in Science, Number 362, pp. 1-13
Natural History Museum of Los Angeles County, 1985

ing an exploratory expedition in 1913. At that time, he con-
sidered the fauna to be early Pleistocene, based on the frag-
mentary mammalian remains (Buwalda, 1914). L.V.
Compton of UCB later published (Compton, 1934) a de-
scription of the five specimens collected by Buwalda and first
noted similarities with the Fossil Lake avifauna. A relatively
large assemblage of birds and mammals was collected in 1952
by H. Winters (1954). The twelve avian taxa from this col-
lection were subsequently described by Howard (1955). The
fauna was considered late Pleistocene in age, and no older
than Illinoian.

Based on fieldwork performed during the middle 1960’s
and a review of existing collections, the Rancholabrean age
assemblage from Lake Manix was designated the Camp Cady
local fauna by G. Jefferson (1968). Initial radiocarbon dating
by various institutions indicated the lake beds were Wiscon-
sinan in age (Bassett and Jefferson, 1971).

Fossil vertebrates from Lake Manix are presently housed
in five separate institutional collections: the Natural History
Museum of Los Angeles County, Vertebrate Paleontology
Section (LACM); San Bernardino County Museum, Earth
Science Collection (SBCM); University of California, Berke-
ley, Museum of Paleontology (UCMP); the University of
California, Riverside, Earth Sciences Department (UCR); and
the United States Geological Survey, Denver, Colorado.

AGE AND STRATIGRAPHY

The Lake Manix avifauna ranges in age from greater than
350 Kyr to about 35 Kyr BP. The stratigraphic section is
well dated by C-14 (carbon- 14) (Bassett and Jefferson, 1971;
Marcus, pers. comm., 1984) and U/Th (uranium-thorium
equilibrium) radiometric techniques (Bischoff, pers. comm.,
1982), and tephrochronologic correlation (Sama-Wojcicki,
1980). Most taxa and the majority of specimens fall between
200 and about 300 Kyr BP (Table 2).

1. George C. Page Museum of La Brea Discoveries, 5801
Wilshire Boulevard, Los Angeles, California 90036.

ISSN 0459-8113

Table 1. Geographic distribution of taxa. Abbreviations: FLO =
Fossil Lake, Oregon; CLC = China Lake, California; RLB = Qua-
ternary Rancho La Brea, California; MKC = McKitterick, Califor-
nia; CAC = Carpinteria, California; sf = subfamily; g = genus; s =

species; ss = subspecies. Extinct taxon = *. Data from: Fortsch, 1978;
Howard, 1946, 1962; Miller, 1925, 1931, 1935; Miller and DeMay,
1942; Stock, 1953.

Lake Manix taxa

FLO

CLC

RLB

MKC CAC

Gavia cf. G. arctica

Podiceps cf. P. nigricollis

s

g

g

Aechmophorus occidentalis

s

g

Pelecanus aff.

P. erythrorhynchos

s

Phalacrocorax auritus

s

g

g

P. macropus*

s

Ciconia maltha*

s

s s

Phoenicopterus minutus*

P. copei*

s

Cygnus cf. C. columbianus

g

g

s

s s

Branta canadensis

s

g

s

s

Anas cf. A. crecca

s

g

s

s

A. cf. A. platyrhynchos

s

g

s

s s

Aythya sp.

g

s

g

Mergus cf. M. merganser

s

Oxyura jamaicensis

s

g

s

Haliaeetus leucocephalus

s

g

s

s s

Aquila chrysaetos

s

g

s

s s

Fulica americana cf.

F. a. shufeldti*

ss

g

s

cf Grus

g

g

g

cf. Actitis

Phalaropodinae

sf

Larus cf. L. oregonus*

s

L. sp.

g

Bubo virginianus

s

s

s s

Lacustrine, fluvial, and alluvial fan deposits of the Manix
Formation (Jefferson, 1968; Jefferson et al., 1982), are well
exposed in bluffs along the Mojave River, 32 km (kilometers)
east of Barstow, San Bernardino County, California. Here,
39 m (meters) of exposed sediment were deposited in a large
freshwater lake that occupied approximately 402 square km,
including the present Coyote and Troy playa lake basins and
Afton Canyon. The horizontal distribution of laterally equiv-
alent sedimentary facies reflects fluctuations in the deposi-
tional system. Fluvial and lacustrine deposits interfinger in
a transgressive/regressive sequence in response to Pleistocene
climatic changes.

Buwalda (1914) recognized a lower and an upper set of
lacustrine deposits within the basin. He attributed their pres-
ence to climatic change and/or tectonic activity.

Winters (1954) described eighteen distinct sedimentary
units in the stratigraphic section, numbered from the top of
the section downward (see columnar section in Jefferson et

Table 2. Stratigraphic/chronologic range of taxa in the Manix
Formation. Abbreviations: m TS = meters above base type section,
Kyr = approximate thousand years. Stratigraphic occurrence = X.
Extinct taxon = *. Data from: Howard, 1955; Jefferson, 1968; Bas-
sett and Jefferson, 1971; Bischoff, pers. comm., 1982.

m TS 9 11 27 30 36

Taxon Kyr BP 350+ 300 200 100 20

Gavia cf. G. arctica

Podiceps cf. P. nigricollis

Aechmophorus occidentalis

Pelecanus aff.

P. erythrorhynchos

Phalacrocorax auritus

P. macropus*

Ciconia maltha*

Phoenicopterus nunutus*

P. copei*

Cygnus cf. C. columbianus
Branta canadensis
Anas cf. A. crecca
A. cf. A. platyrhynchos
Aythya sp.

Mergus cf. M. merganser

Oxyura jamaicensis

Haliaeetus leucocephalus

Aquila chrysaetos

Fulica americana cf.

F. a. schufeldti*

cf. Grus

cf. Actitis

Phalaropodinae

Larus cf. L. oregonus*

L. sp.

Bubo virginianus

X

X X

xx-x-x-x— x-xx

XX

x X-X— X. . .

. . x X . . .

x X. . .

XX

X

X

X-X

X

X

X

X

XX

x X. . .

X

X

X

X

X X

X

al., 1982). Following Buwalda (1914) and Blackwelder and
Ellsworth (1936), Winters recognized two major, climatically
controlled lacustrine phases. He considered the older phase
(units 16 through 14) to be Illinoian in age and the younger
phase (units 13 through 1) temporally correlative with the
Tahoe glaciation. The stratigraphic ranges of avian taxa re-
ported by Howard (1955) are described with reference to
Winters’ numbered units.

Jefferson (1968) divided the Manix Formation into four

2 Contributions in Science, Number 362

Jefferson: Lake Manix Avifauna

members on the basis of lithologically distinctive lacustrine,
fluvial, and lateral fluvial and alluvial facies within the basin.
The lowest unit, member A, is composed of unfossiliferous
alluvial conglomerates (Winters’ unit 18). These rocks are
poorly exposed in the type stratigraphic section (Winters,
1954; Jefferson, 1968; Jefferson et ah, 1982) (located in the
NE XU, SW xk of section 10, T 10 N, R 4 E, U.S.G.S. 15
minute Newberry, California Quadrangle, 1955). The basal
1 1 m of well-exposed deposits in the type section, member
B, mainly consists of fluvial sands and gravels (Winters’ unit
1 7). A U/Th date of 350+ Kyr was obtained from a fragment
of a small Equus sp. humerus located 9 m above the base of
these deposits (Bischoff, pers. comm., 1982). The middle 16
m of exposed section, the lower part of member C, is com-
posed primarily of lacustrine silts and clays. The base of
member C is estimated to be about 290 Kyr BP. The lower
part of member C is approximately equivalent to Buwalda’s
(1914) lower lake and units 16 through 14 of Winters (1954).
The upper 12 m of the Manix Formation, the upper part of
member C and member D, consists of lacustrine silts and
clays overlain by fluvial sands. These deposits are roughly
equivalent to the upper lake beds of Buwalda (1914) and
units 1 3 through 1 of Winters ( 1954). They range in age from
about 200 to 19 Kyr (Bassett and Jefferson, 1971; Sama-
Wojcicki, 1980; Bischoff, pers. comm., 1982; Marcus, pers.
comm., 1984).

All but two avian taxa (Table 2) are known from the basal
1 m of member C (Jefferson, 1968), which consists of very
nearshore silts and sands, and one half of the recorded taxa
(13 of 26) are restricted to this level. This horizon corre-
sponds to unit 16 of Winters (Winters, 1954; Howard, 1955)
and to the base of the lower lake of Buwalda (1914) and
others (Blackwelder and Ellsworth, 1936).

Generally, the more abundant taxa are longer ranging and
exhibit a more continuous stratigraphic record (Table 2 and
Table 3). The stratigraphic distribution of all taxa is restricted
to lacustrine sedimentary facies. Considering the relatively
small sample size for any single taxon, it is probable that
observed biostratigraphic ranges within the Lake Manix sec-
tion are determined by taphonomic factors and do not rep-
resent the temporal range of any taxon.

Other avifauna of similar taxonomic composition (Table
1) are Wisconsinan in age. The Manix avifauna shares most
taxa with the assemblages from Fossil Lake, Oregon and
China Lake, California. Material from Fossil Lake is strati-
graphically associated with a C- 1 4 date of 29 Kyr BP (Allison,
1966). A C-14 date of 18 Kyr (Fortsch, 1978) and a U/Th
date of 42 Kyr (Davis et al., 1981) are associated with the
avifauna from China Lake. The oldest C-14 dates from both
McKittrick (Berger and Libby, 1966) and Rancho La Brea
(Marcus and Berger, 1984) are about 38 Kyr.

SYSTEMATIC DESCRIPTIONS

Most fossil specimens assigned to extant species are mor-
phologically indistinguishable from the modem birds. The
identifications are based primarily on the extensive recent
osteological collections at LACM, and all measurements are
from LACM specimens, unless otherwise noted.

Table 3. Faunal composition. Abbreviations: Extinct taxon = *.

Taxon

Identified

specimens

Relative

percentage

Gavia cf. G. arctica

i

0.7

Podiceps cf. P. nigricollis

4

2.8

Aechmophorus occidenlalis

41

29.5

Pelecanus aff. erythrorhynchos

12

8.6

Phalacrocorax auritus

15

10.8

P. macropus*

2

1.4

Ciconia maltha*

6

4.3

Phoenicopterus minutus*

14

10.1

P. copei*

4

2.8

Cygnus cf. C. columbianus

3

2.1

Branta canadensis

1 1

7.9

Anas cf. A. crecca

1

0.7

A. cf. A. platyrhynchos

2

1.4

Aythya sp.

4

2.8

Mergus cf. M. merganser

2

1.4

Oxyura jamaicensis

3

2.1

Haliaeetus leucocephalus

2

1.4

Aquila chrysaetos
Fulica americana cf

2

1.4

F. a. shufeldti*

1

0.7

cf. Grits

1

0.7

cf. Actitis

1

0.7

Phalaropodinae

1

0.7

Larus cf. L. oregonus*

3

2.1

L. sp.

2

1.4

Bubo virginianus

i

0.7

Total

139

99.5

The use of trinominal names for extinct and extant sub-
species follows Howard (1946). Subspecific names have been
employed by Howard (1946) to distinguish morphologically
and/or proportionally distinct fossil forms from modem
species of the same size range. She considers the extinct sub-
species chronoclinal ancestors to the modem forms. Al-
though the application of subspecific names is noted in the
discussions, they are not applied to fossil populations that
may be larger or smaller than the modem species or fall
within the range of modem subspecies.

Class Aves
Order Gaviiformes
Family Gaviidae
Genus Gavia Foster, 1788

Gavia cf. G. arctica (Linnaeus, 1758)
(Arctic Loon)

REFERRED MATERIAL. SBCM A 500-1506, left and
right angular and dentary.

Contributions in Science, Number 362

Jefferson: Lake Manix Avifauna 3

DISCUSSION. The fragmentary mandible, SBCM A 500-
1 506, compares favorably with modem specimens of G. arc-
tica. It is more slender and not as deep as in G. immer
(Common Loon). Although incomplete, the dorsal margin
of the dentary is straight in lateral view as in G. arctica, not
concave as in G. stellata (Red-throated Loon).

Order Podicipediformes
Family Podicipedidae
Genus Podiceps Brehm, 1831

Podiceps cf. P. nigricollis Brehm, 1831
(Eared Grebe)

REFERRED MATERIAL. UCR 10555, sternum; UCR
10509 and UCR 10576, distal ends of left humeri; UCR
10556, distal right tibiotarsus.

DISCUSSION. Both distal left humeri, UCR 10509 and
10576, are morphologically indistinguishable from modem
specimens of Podiceps. The humeri are distinctly smaller
than the extinct species P. parvus (Shufeldt, 1913) from Fossil
Lake, Oregon.

P. nigricollis and P. auritus (Homed Grebe) are similar in
size and osteologically inseparable. The width of the distal
end of the humerus of P. auritus is generally larger, but over-
laps that of P. nigricollis. A sample of seven humeri of P.
auritus (LACM Ornithology 86335, 86336, 86337, 86339,
86340, and 86341) have a mean width of 7.5 mm (milli-
meters) and the mean of six P. nigricollis (LACM Ornithology
1691, 86329, 86330, 86331, 86332, 86333, and 86334) is
6.9 mm. UCR 10509 and 10576 are closest in size to P.
nigricollis, measuring 7.2 mm and 6.8 mm in width, respec-
tively.

Genus Aechmophorus Coues, 1862

Aechmophorus occidentalis (Lawrence, 1858)
Western Grebe

REFERRED MATERIAL. LACM 2457, partial skeleton
including left and right femora, tibiotarsi, tarsometartarsi,
and other elements (Howard, 1955); LACM 123458, nearly
complete postcranial skeleton; UCR 10542, maxilla, man-
dible, and 4th cervical vertebra; LACM 2466, cervical ver-
tebra (Howard, 1955); LACM 1 12445, cervical vertebra; UCR
14546, cervical vertebra and first tarsal phalanx; LACM
112459, costal margin of sternum; LACM 112414, synsa-
crum; LACM 2465, right coracoid (Howard, 1955); UCR
10538 and 10613, right coracoids; UCR 10882, proximal
right coracoid; LACM 112401, distal left humerus; UCR
10553, proximal left humerus; LACM 2469, right humerus
(Howard, 1955); LACM 123443, proximal left humerus; UCR
14580, 10615, and 10616, right humeri; UCR 10893, prox-
imal right humerus; LACM 1515, fragment humerus; UCR
106 12 and 10874, left femora; UCR 10602, distal left femur;
LACM 123448, proximal right femur; UCR 106 10 and 10611,
right femora; UCR 10552, distal right femur; UCR 13955,

left tibiotarsus and first tarsal phalanx; UCR 10618, right
tibiotarsus; UCR 106 17, distal right tibiotarsus; UCR 10554,
10609 and 10769, proximal left tarsometatarsi; LACM 2458,
proximal right tarsometatarsus (Howard, 1955); UCR 10767,
proximal and distal ends of right tarsometatarsus; UCR
10507, distal right tarsometarsus; UCMP 12859, right tar-
sometatarsus (Compton, 1934); UCR 10614, proximal tar-
sometatarsus; LACM 2459, fragment tarsometatarsus (How-
ard, 1955); UCR 10619, first tarsal phalanx.

DISCUSSION. Miller (191 la) described the extinct sub-
species A. o. lucasi based on the collections from Fossil Lake,
Oregon. Howard (1955:201) described the size of the leg
bones of LACM 2457 as falling “in the zone of overlap of
the Fossil Lake and Recent specimens, although equal to or
greater than the average for A. o. lucasi." Likewise, the skel-
eton, LACM 123458, is osteologically inseparable from the
ranges exhibited by modem specimens of A. occidentalis and
ancestral A. o. lucasi. The femora in LACM 123458 and
LACM 2457 are relatively short compared to the type femur
(UCMP 12605) for A. o. lucasi. The measurements and pro-
portions of the remaining elements, however, are most sim-
ilar to this extinct subspecies. The complete postcranial skel-
eton, LACM 123458, is significant in allowing the description
of the proportions of limb elements relative to isolated ele-
ments. A. o. lucasi is only represented by isolated elements
from Fossil Lake.

In the forelimb of LACM 123458, the lengths of the cor-
acoid, humerus and ulna (Table 4) are very close to the means
of A. o. lucasi from Fossil Lake and modem specimens (How-
ard, 1946). The coracoid falls between the two means, and
the humerus and ulna fall on the mean of A. o. lucasi.

The femur of the Lake Manix specimen, LACM 123458,
measures 42.0 mm in length, which is the same as the small-
est of 63 specimens of A. o. lucasi and well below the mean
of 44.3 mm for eight modem specimens (Howard, 1946).
The type of A. o. lucasi (Miller, 1911a), a femur (UCMP
1 2605), measures 48 mm in length. Measurements of femoral
length in the Fossil Lake sample are greater than the largest
modem specimens and overlap with all but the smallest mod-
em specimens.

The length of the tibiotarsi in LACM 123458, 124 mm
measured to the proximal articular surface, falls very near
the mean of A. o. lucasi reported from Fossil Lake, which is
124.9 mm (Howard, 1946). The total length of this element
ranges from 135 to 147 mm in seven modem specimens
(Gilbert etal., 1981) compared to 146 mm for LACM 123458.
The widths of the proximal and distal ends of the tibiotarsi
measure 1 1.3 mm and 13.3 mm, respectively, compared to
a range of 1 0 to 14 mm for the proximal end of seven modem
specimens (Gilbert et al., 1981).

Tarsometatarsal measurements and ratios (Table 4) of
LACM 123458 are essentially identical to those for A. o.
lucasi listed by Howard (1946, 1955). As with the femora
from Fossil Lake, tarsometatarsi of A. o. lucasi are generally
longer than the modem species and heavier-shafted with a
narrow proximal end (Howard, 1947).

Where comparable measurements are possible, the size
and proportions of isolated skeletal elements from Lake Ma-

4 Contributions in Science, Number 362

Jefferson: Lake Manix Avifauna

Table 4. Measurements for Aechmophorus occidenlalis, LACM
123458. Means of comparable measurements for Recent specimens
are from Howard (1946). Abbreviations: M = mean of Recent spec-
imens; prox. = proximal; dist. = distal; max. = maximum; dia. = di-
ameter; int. = internal; est. = estimated plus or minus 1 mm; intcot.
= intercotylar tubercle.

LACM

123458

M

Coracoid

Length

44.8

44.3

Humerus

Length

1 18.0

1 16.1

Prox. width

19.4

Dist. width

1 1.6

Mid-shaft max. dia.

6.4

Radius

Length

103.4

Prox. max. dia.

4.8

Dist. max. dia.

5.4

Mid-shaft max. dia.

3.3

Ulna

Length

107.1

106.1

Prox. width

8.8

Dist. width

6.0

Mid-shaft dia.

5.3

Femur

Int. length

42.0

44.3

Max. length

46.1

Width dist. condyles

15.!

Tibiotarsus

Length

146 est.

Length to articular surface

124 est.

119.0

Width prox. articular surface

1 1.3

Width dist. condyles

13.3

Tarsometatarsus

Length

77.6

74.5

Shaft width

4.2

Shaft max. depth

7.5

Prox. width

13.7

Width intcot.

5.2

Shaft width/length

5.4%

Shaft max. depth/length

9.7%

Prox. width/length

17.6%

Width intcot. /breadth

123.8%

nix are more similar to LACM 1 23458 than to A. occidentalis
and A. o. lucasi. Coracoids, UCR 10538 and 10613, are
greater than 43 (estimated) and 46.0 mm in length, and fall
between the means of the two taxa. Femora UCR 10610,
10611, and 10612, which measure approximately 44 (esti-

mated), 42.6 and 46.6 mm in length, are relatively small.
The breadth of the intercotylar tubercle relative to the width
of the shaft in tarsometatarsi UCR 10609 and 10767, is 128
and 142 mm (mean 135). This falls below the range for the
Recent form (146-170 mm, mean 1 54) and close to the mean
(140) of A. o. lucasi (Howard, 1946, 1955).

A. o. lucasi is defined on the basis of a femur larger than
the largest Lake Manix specimen. Only one isolated femur
from Fossil Lake is as small as those from Lake Manix.
LACM 2457 and LACM 123458 have small femora, and
although close to A. o. lucasi in all other skeletal dimensions,
cannot be assigned to the late Pleistocene extinct subspecies
from Fossil Lake, Oregon. The assemblage from Fossil Lake
is significantly younger than the Lake Manix assemblage which
may account for these minor proportional differences.

Order Pelecaniformes

Family Pelecanidae

Genus P elec anus Linnaeus, 1758

P elec anus aff. P. erythrorhynchos Gmelin, 1789
(American White Pelican)

REFERRED MATERIAL. UCR 14724, anterior ster-
num; UCR 10633, proximal left scapula; LACM 2460, left
coracoid (Howard, 1955); UCR 20991, proximal right cor-
acoid; LACM 123445, right humerus; UCR 1 5669, proximal
humerus fragment; LACM 2462, proximal right radius
(Howard, 1955); LACM 123447, proximal left carpometa-
carpus; LACM 2461, distal right femur (Howard, 1955);
UCMP 21855, left femur (Compton, 1934); UCMP 12857,
first tarsal phalanx left second digit (Compton, 1934); UCMP
12858, first tarsal phalanx, left third digit (Compton, 1934).

DISCUSSION. As noted by Howard (1955) and con-
firmed by additional specimens, the Lake Manix pelican is
morphologically the same, but larger than the extant Amer-
ican White Pelican. No fossils are smaller than the largest
modem specimens examined. The complete right humerus,
LACM 123445, measures 336 mm in length compared to
five modem specimens which range from 284 to 321 mm
(Gilbert et al., 1981).

The Lake Manix material apparently represents a robust
population of P. erythrorhynchos. Although criteria such as
size alone have been used previously by others to define fossil
avian subspecies or chronoclinal races, I do not concur with
the practice. Proportional differences in limb measurements
between the modem and fossil specimens cannot be dem-
onstrated on the basis of the small Lake Manix sample.

Family Phalacrocoracidae
Genus Phalacrocorax Brisson, 1760

Phalacrocorax auritus (Lesson, 1831)
Double-crested Comorant

REFERRED MATERIAL. LACM 123457, nearly com-
plete postcranial skeleton; UCR 10577, premaxilla fragment;

Contributions in Science, Number 362

Jefferson: Lake Manix Avifauna 5

Table 5. Measurements for Phalacrocorax auritus, LACM 123457.
Abbreviations: M = mean of four large Recent specimens (LACM
Ornithology 100734, 100831, 100832, and 101213); prox. = proxi-
mal; dist. = distal; dia. = diameter; max. = maximum; est. = esti-
mated plus or minus 1 mm; proc. = process; int. = internal.

LACM

123457

M

Scapula

Length

89.2

85.0

Prox. width

18.9

18.0

Humerus

Length

161.2

154.8

Prox. width

25.0

22.9

Dist. width

17.7

16.6

Mid-shaft dia.

8.8

8.2

Radius

Length

168.1

160.0

Prox. max. dia.

8.9

8.6

Dist. max. dia.

9.3

9.5

Mid-shaft dia.

4.5

4.6

Ulna

Length

168 est.

164.5

Prox. width

13.4

12.9

Dist. width

10.0

10.1

Mid-shaft dia.

6.6

6.3

Carpometacarpus

External length

77.4

74.9

Prox. depth

15.0

13.9

Height proc. metacarpal 1

11.7

11.1

Femur

Int. length

61.7

57.9

Max. length

64.0

60.4

Dist. width

18.9

16.9

Tibiotarsus

Length to prox. articular surface

113.7

107.8

Width dist. condyles

14.1

13.1

UCR 10635, left quadrate; UCR 15667, right scapula; UCR
10637, left coracoid; SBCM A 1768-1, distal right coracoid
and proximal right femur fragment; UCR 10627, left hu-
merus; UCR 14684, distal right humerus; LACM 2468, distal
left ulna; UCR 10557, proximal right carpometacarpus; UCR
10623, distal right carpometacarpus; UCR 14666, proximal
and distal ends of tibiotarsus; SBCM A 500-1500, proximal
left tarsometatarsus; UCR 10630, distal right tarsometatar-
sus; UCR 14547, first tarsal phalanx.

DISCUSSION. Lake Manix material compares very fa-
vorably with the largest modem specimens of P. a. alboci-
liatus (Table 5) from the Pacific coast. Measurements of the
principal limb elements of LACM 123457 (Table 5) are dis-

tinctly smaller than those given for extinct P. macropus (Cope,
1878) (Howard, 1946), and are slightly larger than the largest
modem specimens examined or listed by Gilbert et al. ( 1 98 1 ).
Large size is also evident in all isolated limb elements.

Howard (1932) placed subgeneric value on the ratio of the
distance from the anterior intermuscular line to the internal
edge of the coracoid compared to the length of the coracoid
measured to the internal sternal lip. The subgenus Phalacro-
corax (P. auritus, P. carbo, and P. olivaceus) has a ratio of
15 percent, compared to 19-21 percent in the subgenus
Compsohalieus (P. pencillatus) and 21-22 percent in the sub-
genus Urile (P. pelagicus and P. perspicillatus) (Howard, 1 946).
P. macropus yields a ratio of 17-19 percent. This ratio for
LACM 123457 (11.7 mm, 71.9 mm) is 16.3 percent. The
isolated left coracoid, UCR 10637, measures 65.2 and 1 1.0
mm and yields a ratio of 16.9 percent.

Measurements of isolated elements are also larger than the
largest P. auritus examined, although smaller than P. mac-
ropus. Width of the distal end of humerus UCR 10627 is
1 7.5 mm and mid-shaft diameter is 9.8 mm. Distal humerus,
UCR 14684 measures 15.6 mm in width. In UCR 14666,
the width of the distal end of the tibiotarsus measured across
the proximal edge of the supratendinal bridge is 13.1 mm.
The proximal end of the specimen is 14.2 mm in width. The
left proximal tarsometatarsus, SBCM A 500-1500, measures
14.0 mm in width, 19.1 mm in depth, and has a mid-shaft
width of 7.3 mm and depth of 7.1 mm. The ridge supporting
the external cotyle in this specimen is thickened and more
prominent than in modem specimens.

The Lake Manix material apparently represents a robust
population of P. auritus larger than the living west coast form
(Table 5). No intermediate-sized individuals are present in
the sample. Relative limb proportions of LACM 123457 and
in modem specimens are essentially identical.

Phalacrocorax macropus Cope, 1878

REFERRED MATERIAL. SBCM A 500-1382, premax-
illa; LACM 123442, proximal right humerus.

DISCUSSION. The nearly complete premaxilla, SBCM
A 500-1382 (Fig. 1), is damaged along the proximal edge and
is missing about 2-3 mm of bone tissue. Its dorsal margin
in lateral view is more concave than in P. auritus. The spec-
imen is more robust than the premaxilla of either P. auritus
or P. pencillatus relative to its comparatively short length of
62 mm. It measures 16.8 mm in width at the distal margin
of the nasal aperture and 13.3 mm in dorsal-ventral height,
exceeding the size of both large modem species. These fea-
tures closely conform to Shufeldt’s (1892) description of the
upper mandible of P. macropus from Fossil Lake, Oregon.

Although the proximal ends of the humeri are not known
for P. macropus, the size and thickness of the mid-shaft of
LACM 123442 support its placement here. The humerus
measures 26. 1 mm across the proximal end and has a mid-
shaft diameter of 10.4 mm. A large modem specimen of P.
auritus (LACM Ornithology 100734) measures 24.2 mm in
proximal width, 16.2 mm across the distal end, and has a
mid-shaft diameter of only 8.4 mm. Gilbert et al. (198 1) give

6 Contributions in Science, Number 362

Jefferson: Lake Manix Avifauna

the range for the proximal width of six modem specimens
as 23 to 24 mm. The width of the distal humerus of P.
macropus ranges from 18.6 to 19.8 mm (Howard, 1946).

Order Ciconiiformes
Family Ciconiidae
Genus Ciconia Linnaeus, 1758
Ciconia maltha Miller, 1910

REFERRED MATERIAL. LACM 2463, right humerus
(Howard, 1955); UCR 13637, proximal right femur; UCR
14732, proximal right tibiotarsus; UCR 10629, proximal and
distal right tarsometatarsus; UCR 10628 and 10759, first
tarsal phalanges.

DISCUSSION. All fossil Ciconia material from California
has been referred to C. maltha (Howard, 1942). The Lake
Manix specimens are indistinguishable from equivalent ele-
ments in the Rancho La Brea sample.

Order Phoenicopteriformes
Family Phoenicopteridae
Genus Phoenicopterus Linnaeus, 1758
Phoenicopterus minutus Howard, 1955

REFERRED MATERIAL. LACM 2446, left scapula
(Howard, 1955); SBCM A 500-1507, proximal left scapula;
UCR 14669, right scapula; LACM 2474, right coracoid frag-
ment and scapula (Howard, 1955); UCR 10578, proximal
right coracoid; UCR 10631, left coracoid; UCR 10636, prox-
imal first carpal phalanx; SBCM A 500-1504, proximal left
femur; UCR 14544, proximal right femur; LACM 2445 (type
specimen), right tibiotarsus and proximal tarsometatarsus
(Howard, 1955); LACM 1 12438, distal left tibiotarsus; LACM

2473, proximal left tarsometatarsus (Howard, 1955); UCR
10875, proximal right tarsometarsus.

DISCUSSION. Howard (1955) described P. minutus as a
very small flamingo, smaller than extinct P. stocki (Miller,
1 944) from the Pliocene of Mexico. It differs from P. stocki,
the larger extinct P. copei (Shufeldt, 1892) and the modern
P. chiliensis (Chilean Flamingo), and P. ruber (Greater Fla-
mingo) in minor, but distinctive, morphologic characters.
Howard (1955) provisionally referred a left scapula, LACM
2446, and a right coracoid fragment and scapula, LACM

2474, to this taxon. Comparable additional specimens of
these elements, SBCM A 500-1 507, UCR 14669, 10578, and
10631, now support the assignment of this material to P.
minutus.

Femora were not previously known for P. minutus. SBCM
A 500-1504 and UCR 14544 are closely comparable to, but
distinctly smaller than, modem species of Phoenicopterus as
well as the extinct P. copei (Shufeldt, 1 892). A narrow shallow
groove common to all the species of Phoenicopterus is present

A

B

Figure 1. Pha/acrocorax macropus premaxilla, SBCM A 500-
1382. A, right lateral view. B, dorsal view. Scale bar is 30
mm.

on the lateral margin of the crest in both specimens. The
width of SBCM A 1500-1504 is 16.8 mm. The trochanteric
ridge visible in this specimen swings medially towards the
head of the femur, as in P. ruber. UCR 14544, although
incomplete and abraded along the dorsal edge of the tro-
chanter, measures 16.1 mm in width.

A proximal carpal first phalanx, UCR 10636, although
morphologically comparable, is smaller than specimens of
P. ruber and tentatively referred to P. minutus.

Phoenicopterus copei Shufeldt, 1892

REFERRED MATERIAL. LACM 112410, cervical ver-
tebra; LACM 2448, left tarsometatarsus (Howard, 1955);
UCR 10879, right tarsometarsus and three associated pha-
langes; LACM 123440 and UCR 14693, medial first tarsal
phalanges.

DISCUSSION. Howard (1955) placed the immature left
tarsometatarsus, LACM 2448, in the extinct species P. copei
based on comparison with measurements of the distal con-
dyles of tibiotarsi from Fossil Lake, Oregon. Tarsometatarsi
are not present in the Fossil Lake sample. The widths of the
fossil tarsometatarsi are greater than that of modem species
of Phoenicopterus { Howard, 1946).

Measurements of the right tarsometatarsus, UCR 10879,
are very close to LACM 2448, which measures 328 mm in
length, 19 mm in width across the proximal articular surface,
12 mm in intercotylar width, and 6.8 mm in height of in-
tercotylar tubercle. UCR 10879 measures 333.6 mm in length,
18.6 mm in width across the proximal articular surface, 12.1
mm in width across the intercotylar tubercle, and the height
of the intercotylar tubercle is 6.8 mm (Jefferson, 1968).

Contributions in Science, Number 362

Jefferson: Lake Manix Avifauna 7

The two medial tarsal first phalanges, LACM 123440 and
UCR 14693, are large but otherwise comparable to P. ruber.

Order Anseriformes
Family Anitidae
Genus Cygnus Linnaeus, 1758

Cygnns cf. C. columbianus (Ord, 1815)
(Whistling Swan)

REFERRED MATERIAL. UCR 10634, scapula; UCR
14725, distal left tibiotarsus; UCR 13953, lateral first tarsal
phalanx.

DISCUSSION. The scapula, UCR 10634, is not separable
from modem specimens of C. columbianus. UCR 14725, a
distal left tibiotarsus, measures 18.9 mm in width, which is
slightly smaller but otherwise closely comparable to a sample
of four modem specimens (LACM Ornithology 86547, 86548,
86552, and 86553 that range from 20.5 to 24.7 mm in width).
It is clearly separable from similar-sized specimens of Branta
where the tendinal groove is distinct and extends to the mid-
shaft. The distal margin of the supra-tendinal bridge is con-
cave in Cygnus and straight in Branta. The lateral tarsal
phalanx, UCR 13953, is morphologically comparable to C.
columbianus although more slender.

C. buccinator (Trumpeter Swan) and the extinct species C.
paloregonus (Cope, 1878) are present in the Fossil Lake,
Oregon, fauna. Both taxa exceed the dimensions of C. co-
lumbianus, which is not found at Fossil Lake.

Genus Branta Scapoli, 1769

Branta canadensis Linnaeus, 1758
Canada Goose

REFERRED MATERIAL. UCR 10621, maxilla; LACM
123450 and UCR 10622, left scapula; LACM 123449, left
coracoid; LACM 112409 and UCR 10625, fragment right
coracoid; UCR 10624, right coracoid; UCR 10894, proximal
and distal left humerus; UCR 10626, partial right humerus;
UCR 10620, distal left ulna; LACM 2464, proximal and
distal right femur (Howard, 1955).

DISCUSSION. The Lake Manix specimens are separable
into large and small forms that do not overlap in size. The
larger specimens (UCR 10621, maxilla; LACM 123450 and
UCR 10622, left scapula; LACM 123449, left coracoid;
LACM 112409 and UCR 10625, fragment right coracoid;
UCR 1 0624, right coracoid; UCR 1 0894, proximal and distal
left humerus; LACM 2464, proximal and distal right femur)
are closely comparable in morphology and size to modem
specimens of the largest subspecies of B. canadensis, B. c.
canadensis.

The relatively small distal left ulna, UCR 10620, is iden-
tical in size and morphology to specimens of the Recent B.
c. minima, smallest subspecies of B. canadensis.

Although clearly assignable to Anserini, both ends of the
humerus, UCR 10626, are badly damaged. This specimen is

tentatively considered a small B. canadensis based on an
approximate length of 160 mm.

The largest and smallest (nominal) subspecies of B. can-
adensis are also represented at Fossil Lake, Oregon (Howard,
1955). In this respect, the limited Lake Manix sample appears
to parallel the Fossil Lake avifauna. However, the material
does not warrant subspecific assignment.

Genus Anas Linnaeus, 1758

Anas cf. A. crecca Linnaeus, 1758
(Green-winged Teal)

REFERRED MATERIAL. LACM 112415, right humer-
us.

DISCUSSION. Approximately 1 to 2 mm of bone tissue
have been abraded from the head and entepicondyle of the
right humerus, and the external tuberosity and pectoral at-
tachment are missing. Although the specimen is incomplete,
a total length of 56 mm is comparable with the size of A.
crecca.

Anas cf. A. platyrhynchos Linnaeus, 1758
(Mallard)

REFERRED MATERIAL. UCR 10551, partial right hu-
merus; UCR 13956, proximal right humerus.

DISCUSSION. The end of right humerus UCR 10551
proximal to the bicipital crest is missing. The preserved por-
tion is 82 mm in length. A total estimated length of 1 03—
104 mm is slightly larger than the largest modem A. platy-
rhynchos examined.

The proximal humerus, UCR 13956, is inseparable from
equivalent elements in the Rancho La Brea sample of A.
platyrhynchos. It is distinguished from similar-sized small
members of the Anserini by a more laterally directed median
crest and deeper ligamental furrow.

Genus Aythya Boie, 1822
Aythya sp.

Greater Scaup or Canvasback

REFERRED MATERIAL. UCR 14545, left coracoid;
LACM 2472, right scapula (Howard, 1955); LACM 2475,
distal left humerus (Howard, 1955); SBCM AE 873-2, left
femur.

DISCUSSION. A. marila (Greater Scaup) and A. valisi-
neria (Canvasback) are not easily separated osteologically.
The lack of diagnostic specimens precludes a specific assign-
ment. Howard (1955) tentatively referred LACM 2472 and
2475 to A. valisineria.

Genus Mergus Linnaeus, 1758

Mergus cf. M. merganser Linneaus, 1758
(Common Merganser)

REFERRED MATERIAL. UCR 10895 and 14581, left
coracoids.

8 Contributions in Science, Number 362

Jefferson: Lake Manix Avifauna

DISCUSSION. Of the numerous modern coracoids ex-
amined, those of M. merganser are generally larger than, but
show considerable size overlap with the coracoids of M. ser-
rator (Red-breasted Merganser). The fossil coracoids fall
within the size range of M. merganser and M. serrator. How-
ever, an excavated area posterior to the brachial tuberosity
and antero- ventral to the scapular facet is consistently deeper
in M. merganser compared to M. serrator. Both UCR 10895
and 14581 clearly exhibit this character.

Genus Oxyura Bonaparte, 1828

Oxyura jamaicensis (Gmelin, 1789)

Ruddy Duck

REFERRED MATERIAL. LACM 2476, anterior ster-
num (Howard, 1955); LACM 1 12433, distal left humerus.

DISCUSSION. Lake Manix specimens are indistinguish-
able from modern specimens of Ruddy Duck. Although the
width (8.8 mm) of the distal left humerus, LACM 1 12433,
is close to that of Anas crecca, it is distinguished by having
a larger anterior articular ligamental attachment and straight-
er shaft.

Order Aecipiiri formes
Family Accipitridae
Genus Hciliaeetus Savigny, 1 809

Haliaeetus leucocephalus Linnaeus, 1758
Bald Eagle

REFERRED MATERIAL. UCR 10638, distal right cor-
acoid; UCR 14739, distal humerus.

DISCUSSION. These specimens compare closely with the
Rancho La Brea sample of H. leucocephalus. The lateral
flange on the posterior margin of the coracoid, UCR 10638,
is wide and moderately hooked proximally. This condition
is present on a minority of the Rancho La Brea specimens
and absent from ail available Recent material (Jefferson,
1968). The more laterally positioned scar for M. coracobra-
chialis in UCR 10638 clearly separates it from similar-sized
coracoids of the extinct hawk, Amplibuteo woodward i (Mil-
ler, 1911b).

Genus Aquila Brisson, 1760

Aquila chrysaetos (Linnaeus, 1758)

Golden Eagle

REFERRED MATERIAL. LACM 2470, distal right tibio-
tarsus (Howard, 1955); UCR 10632, fused first and second
tarsal phalanges of left second digit.

DISCUSSION. The Lake Manix specimens are insepa-
rable from the Rancho La Brea sample of Aquila chrysaetos.
AMNH (American Museum of Natural History) 3467B, a
second tarsal phalanx of digit two from Fossil Lake, Oregon,
was first assigned to A. chrysaetos by Shufeldt (1892). This
specimen, which was later referred to Spizaetus pliogryps by

Howard (1946), is more slender than UCR 10632 and spec-
imens of A. chrysaetos from Rancho La Brea. UCR 10632
measures 37.8 mm in length, 14.1 mm in proximal width,
and 12.0 mm in proximal dorso-ventral height.

Order Gruiformes
Family Rallidae
Genus Fulica Gmelin, 1789

Fulica americana Gmelin, 1789
American Coot

Fulica americana cf. shufeldti Brodkorb, 1964

REFERRED MATERIAL. LACM 123446, left ulna.

DISCUSSION. Howard (1946, 1947) considered the rel-
atively small, extinct coot, F. minor (Shufeldt, 1892) from
Fossil Lake, Oregon, to be an ancestral subspecies (chrono-
cline) of the modem form, F. americana, for which she erect-
ed subspecies F. a. minor. Subsequently Wetmore (1956)
elevated “ minor ” to specific level following Shufeldt’s orig-
inal designation. Brodkorb (1964), recognizing that the name
minor was previously occupied in the genus, renamed the
taxon after Shufeldt, calling the form F. shufeldti.

The measurements given by Howard (1946) clearly show
an overlap in the size of the specimens from Fossil Lake,
Oregon, and modern specimens. There is no question that a
single species is represented. Following Howard, the extinct
form is considered a subspecies of F. americana, and is re-
ferable to F. a. shufeldti.

The ancestral subspecies differs from the modem Ameri-
can Coot in having shorter wings and relatively long legs
(Howard, 1946, 1947). Mean length of three Fossil Lake
ulnae is 56.9 mm, compared to 61.1 mm for 39 modem
specimens (Howard, 1946). The Lake Manix ulna, LACM
123446, measures 56.8 mm.

Genus Grus Pallas, 1766
cf. Grus sp.

Crane

REFERRED MATERIAL. LACM 2467, fragmentary dis-
tal right humerus (Howard, 1955).

DISCUSSION. Howard (1955) questionably assigned this
specimen to Grus. No additional crane specimens have been
recovered.

Order Charadriiformes
Family Scolopacidae
Subfamily Scolopacinae
Genus Ac this Illiger, 1811

cf. Actitis sp.

(Sandpiper)

REFERRED MATERIAL. LACM 123451, proximal left
humerus.

Contributions in Science, Number 362

Jefferson: Lake Manix Avifauna 9

DISCUSSION. Various tribes within the subfamily Scol-
opacinae cannot be readily separated. LACM 123451 best
compares with Actitis hypoleucos (Common Sandpiper) in
size and approaches the smallest species of Tringa (sandpi-
pers and yellowlegs). It differs from humeri of Phalaropus
lobatus (Red-necked Phalarope, subfamily Phalaropodinae),
which are almost identical in size, in having a deeper and
more enclosed pneumatic fossa and a more rounded lateral
margin of the bicipital crest.

Subfamily Phalaropodinae
Genus and species indet.

REFERRED MATERIAL. LACM 247 1 , partial left tibio-
tarsus (Howard, 1955).

DISCUSSION. LACM 247 1 , a very small, left distal tibio-
tarsus, was placed within the Phalaropodinae but could not
be confidently assigned to the genus Phalaropus by Howard
(1955). No additional comparable specimens have been re-
covered.

Family Laridae
Genus Larus Linnaeus, 1758
Larus cf. L. oregonus Shufeldt, 1892

REFERRED MATERIAL. LACM 1 23444, associated left
humerus, radius, and carpometacarpus; LACM 123802, left
first carpal phalanx, digit two.

DISCUSSION. The Lake Manix specimen, LACM 1 23444
(Fig. 2), represents a robust, medium-sized gull morpholog-
ically similar to L. canus (Mew Gull) and L. delawarensis
(Ring-billed Gull). It is larger and more robust than the largest
L. canus examined, and smaller than L. delawarensis. Di-
mensions of the limb elements differ from the wing of L.
delawarensis in being shorter and having thicker or stouter
mid-shafts. It is most similar to extinct L. oregonus from
Fossil Lake, Oregon.

The humerus, although slightly smaller, is comparable in
robustness to the type of L. oregonus. The maximum width
of the proximal end of LACM 123444 (the bicipital crest
margin shows slight damage) is 16.8 mm, compared to 17.7
mm in the type specimen, AMNH 3494 (measurements from
cast). Minimum mid-shaft breadth is 5.8 mm, compared to
6.6 mm in the type, and the breadth of the shaft immediately
distal to the base of the bicipital crest in LACM 123444 is
8.8 mm, versus 9.1 mm.

Howard (1946:186), in describing AMNH 3494, stated
that “the depression below the head anconally . . . bounded
by the sharp apex of the shaft on one side and the median
crest on the other, tends to narrow almost to a point.” This
condition in LACM 123444 is similar to that in L. canus.
The depression in L. delawarensis is more rounded, inter-
mediate between L. oregonus and the wide depression in L.
californicus. In this character, LACM 123444 is less pointed

Figure 2. Larus cf. L. oregonus associated left wing ele-
ments, LACM 123444. A, anconal view of humerus. B, an-
conal view of radius. C, external view of carpometacarpus.
Scale bar is 30 mm.

than AMNH 3494, not as wide or rounded as in L. dela-
warensis, and most similar to the largest specimens of L.
canus, which exhibit the most rounded condition in this
species.

Radii and carpometacarpi of L. oregonus are not known
from Fossil Lake. The Lake Manix radius, although almost
a centimeter shorter than the radii of L. delawarensis, mea-
sures 102. 1 mm in length and is markedly stouter. Maximum
thickness of the proximal portion of the shaft is 3.6 mm,
much greater than in L. canus (LACM Ornithology 87200,
3.2 mm; 87201, 3.2 mm) or L. delawarensis (LACM Orni-
thology 87204, 3.1 mm; 87205, 3.2 mm; 87206, 2.9 mm;
87207, 3.1 mm). The bicipital tubercle is more pronounced
and distally elongated in LACM 123444, and the groove
bounded by the interosseous crest is longer and deeper in
palmar view.

The carpometacarpus is morphologically similar to that of
L. canus and L. delawarensis, but it is shorter and exhibits
a considerably thickened main shaft. It measures 53.3 mm
in length. LACM 123802, a first carpal phalanx, articulates

30 Contributions in Science, Number 362

Jefferson: Lake Manix Avifauna

almost exactly with the carpometacarpus of LACM 123444,
and measures 24.8 mm in length.

Larus sp.

Gull

REFERRED MATERIAL. LACM 123803, left proximal
radius, coracoid, distal ulna, and proximal and distal car-
pometacarpus; SBCM A 500-1508, proximal portion of left
carpometacarpus; SBCM A 500-1505, distal left carpometa-
carpus.

DISCUSSION. The specimens represent a large gull of
the size of L. argentatus (Herring Gull) or L. occidentalis
(Western Gull), but larger than L. californicus (California
Gull). Larger specimens of L. occidentalis and L. californicus
exhibit considerable overlap in size with smaller specimens
ofL. argentatus. LACM 123803 and SBCM A 500-1508 fall
within this range. However, SBCM A 500-1505 is slightly
larger than the largest L. argentatus examined. The fossil
specimens cannot be distinguished from larger species of the
genus Larus.

Remains of gulls are extremely rare in Quaternary marine
deposits of western North America (Miller, 1924; Brodkorb,
1967). Although abundant and varied at Fossil Lake, Oregon
(Howard, 1946), fossil specimens representing the Laridae
previously were not known from Pleistocene asphalt or other
terrestrial deposits in California (Miller and DeMay, 1942;
Howard, pers. comm., 1983). The record from late Pleisto-
cene marine deposits includes a single specimen of Larus
glaucescens from the upper San Pedro Sand, California (Mil-
ler, 1930) and a second specimen from New Port Bay Mesa,
California (Howard, 1 949). The distribution of Larus at Ran-
cho La Brea (Howard, 1936) and in Alaska (Brodkorb, 1967)
is restricted to the Holocene. The only known Tertiary spec-
imen is from the Pliocene San Diego Formation of southern
California (R. Ceruti, pers. comm., 1983). Although samples
are small, taphonomic or preservational biases do not seem
to fully account for the very poor representation of gulls at
Rancho La Brea or in late Pleistocene coastal marine deposits
that yield numerous remains of other, similar-sized avian
taxa (Howard, 1949). The inland late Pleistocene record of
gulls appears to contrast with their appearance in Holocene
coastal communities. These records suggest that a shift in
the distribution or population density of gulls may have oc-
curred in response to the disappearance of most inland la-
custrine habitats at the end of the Pleistocene.

Order Strigiformes
Family Strigidae
Genus Bubo Dumeril, 1806

Bubo virginianus (Gmelin, 1788)

Great Homed Owl

REFERRED MATERIAL. UCR 10514, distal left tibio-
tarsus.

DISCUSSION. The width of the distal condyles on tibio-
tarsus UCR 1 05 1 4 is 13.1 mm, identical to the mean of fifty
Rancho La Brea specimens which range from 12.2 to 13.4
mm. A sample of twelve Recent specimens range between
1 1.9 and 13.4 mm (Howard, pers. comm., 1967). The large
extinct owl. Bubo sinclairi (Miller, 1911c), overlaps the size
of the largest Rancho La Brea specimens, exceeding UCR
10514.

BIOGEOGRAPHY AND PALEOENYIRONMENT

All of the extant species represented in the Lake Manix avi-
fauna are at least seasonally present in southern California.
Most taxa are found along the California coast during the
winter or are winter visitors on inland lakes, such as the
Salton Sea, or along the Colorado River. Pelecanus is a sum-
mer visitor most common at the Salton Sea. Only Oxyura
and Fulica are wide ranging throughout the year. Cygnus is
rare in southern California and found in winter on inland
lakes or reservoirs to the north (Cogswell and Christman,
1977; Garrett and Dunn, 1981).

All extant, migratory species leave southern California in
the spring. They travel northward along the coast or follow
inland portions of the north-south Pacific Coast flyway. Dur-
ing Pleistocene pluvial periods, this inland route would have
been over the lakes of the Mojave Desert, the lakes east of
the Sierra Nevada Mountains including China Lake, the
western part of Lake Lahontan, and the lakes of southeastern
Oregon, including Fossil Lake (Snyder et al., 1964).

Two-thirds of the extant taxa (9 of 1 7), represented by 80
percent of the fossil specimens (Table 3), presently prefer, or
feed exclusively on, small fish (Cogswell and Christman, 1 977):
Gavia arctica, Podiceps nigricollis, Aechmophorus occiden-
talis, Pelecanus erythrorhynchos, Phalacrocorax auritus,
Mergus merganser, Aquila chrysaetos, Haliaeetus leucoceph-
a/us, and Larus spp. Abundant fossil remains of the small
Tui (Mojave) Chub, Gila bicolor mojavensis, are present in
the lacustrine deposits (Jefferson, 1 968). No other fishes have
been reported.

Most of the remaining taxa feed on a variety of water plants
and freshwater invertebrates (Cogswell and Christman, 1 977):
Cygnus columbianus. Anas crecca, A. platyrhynchos, Oxyura
jamaicensis, Fulica americana, Act it is sp., and Brant a can-
adensis. Aquila is mainly a scavenger, and Bubo virginianus
primarily feeds on small mammals.

The Lake Manix assemblage samples a complex of fresh-
water lake and lake margin habitats. Judging from food pref-
erences, procurement methods, and nesting habits (Cogswell
and Christman, 1977) of the extant forms represented, open
water, sandy beach flats, and extensive reedy marshlands
must have been persistent lacustrine features. An extensive
lacustrine environment is confirmed by lithostratigraphy and
reconstructions of the depositional environments (Jefferson,
1968).

ACKNOWLEDGMENTS

The encouragement and continued advice of H. Howard are
sincerely appreciated. I thank colleagues at the Natural His-

Contributions in Science, Number 362

Jefferson: Lake Manix Avifauna 11

tory Museum of Los Angeles County who offered valuable
comments and critically reviewed the manuscript. R. Schrei-
ber and K. Garrett in the Ornithology Section of the Museum
provided the modem, comparative osteological specimens
essential to this study. I also wish to thank R.E. Reynolds of
the San Bernardino County Museum and M. Woodbume
and W. Daily of the University of California at Riverside for
the loan of specimens under their care. J. Brinkerhoff and G.
Drewes donated fossil gull specimens (LACM 123802 and
123803) important to this study.

LITERATURE CITED

Allison, I.S. 1966. Fossil Lake, Oregon: Its geology and
fossil faunas. Oregon State Univ., Studies in Geology 9:
1-48.

Bassett, A. M., and G.T. Jefferson. 1971. Radiocarbon dates
of Manix Lake, central Mojave Desert, California. Geo-
logical Soc. of America Spec. Pap., Abs. to Meetings
3(2):79.

Berger, R., and W.F. Libby. 1966. UCLA radiocarbon dates
V. Radiocarbon 8:492.

Blackwelder, E., and E.W. Ellsworth. 1936. Pleistocene lakes
of the Afton basin, California. American Jour. Sci. 5th
Ser. 31:453-463.

Brodkorb, P. 1 964. A new name for Fulica minor Shufeldt.
Quart. Jour. Fla. Acad. Sci. 27(3): 186.

. 1967. Catalog of fossil birds: Part 3 (Ralliformes,

Ichthyomithiformes, Charadriiformes). Florida State
Mus. Bull. Biological Sciences 2(3).

Buwalda, J.P. 1914. Pleistocene beds at Manix in the east-
ern Mojave Desert region. Bull. Dept. Geology Univ.
Calif., Berkeley 7(24):443-464.

Cogswell, H.L., and G. Christman. 1977. Water birds of
California. Univ. Cal. Press, Berkeley, Calif. Nat. Hist.
Guides 40:v + 399.

Compton, L.V. 1934. Fossil bird remains from the Manix
Lake deposits of California. Condor 36:166-168.

Cope, E.D. 1878. Description of new Vertebrata from the
Upper Tertiary and Dakota formations. U S. Geol. and
Geog. Surv. Terr. Bull. 4:379-396.

Davis, E.L., G.T. Jefferson, and C. McKinney. 1981. Man-
made flakes with a dated mammoth tooth at China Lake,
California. Anthropological Jour, of Canada 1 9(2):2— 7.
Fortsch, D.E. 1978. The Lake China Rancholabrean fau-
nule. In The ancient Californians: Rancholabrean hunt-
ers of the Mojave Lakes Country, eds. E.L. Davis, 173—
176. Nat. Hist. Mus. of Los Angeles Sci. Ser., 29.
Garrett, K., and J. Dunn. 1981. The birds of southern Cal-
ifornia status and distribution. Los Angeles Audubon
Soc., viii + 408.

Gilbert, B.M., L.D. Martin, and H.G. Savage. 1981. Avian
osteology’. B. Miles Gilbert Publisher, i + 252.

Howard, H. 1932. A new species of cormorant from Plio-
cene deposits near Santa Barbara, California. Condor
34:118-120.

. 1936. Further studies upon the birds of the Pleis-
tocene of Rancho La Brea, California. Condor 38:32-
36.

. 1942. A review of the American fossil storks. Car-
negie Inst. Wash. Publ. Contrib. to Paleont. 530:187-
203.

. 1 946. A review of the Pleistocene birds from Fossil

Lake, Oregon. Carnegie Inst. Wash. Publ. 551:141-195.

. 1 947. A preliminary survey of trends in avian evo-
lution from Pleistocene to Recent time. Condor 49(1):
10-13.

. 1949. Avian fossils from the marine Pleistocene of

southern California. Condor 51(1 ):20— 28.

. 1955. Fossil birds from Manix Lake, California.

U.S. Geological Survey Prof. Pap. 264-J: 199-205.

. 1962. A comparison of avian assemblages from

individual pits at Rancho La Brea, California. Los An-
geles Co. Mus. Contrib. Sci. 58:1-24.

Jefferson, G.T. 1968. The Camp Cady local fauna from
Pleistocene Lake Manix, Mojave Desert, California.
Dept. Geol. unpubl. masters thesis, Univ. Cal., River-
side, vii -I- 130.

Jefferson, G.T., J.R. Keaton, and P. Hamilton. 1982. Ma-
nix Lake and the Manix Fault field trip guide. Quarterly
San Bernardino Co. Mus. Assoc. 29(3-4): 1-47.

Marcus, L.F., and R. Berger. 1984. The significance of ra-
diocarbon dates for Rancho La Brea. In Quaternary ex-
tinctions a prehistoric revolution, ed. P.S. Martin, and
G. Klein, 159-183. Univ. Arizona Press.

Miller, L. H. 1911a. Additions to the avifauna of the Pleis-
tocene deposits at Fossil Lake, Oregon. Univ. Cal. Publ.
Dept. Geol. Sci. 6:79-87.

. 1911b. A series of eagle tarsi from the Pleistocene

of Rancho La Brea. Univ. Cal. Publ. Dept. Geol. Sci. 6:
305-316.

. 1911c. Avifauna of Pleistocene cave deposits of

California. Univ. Cal. Publ. Dept. Geol. Sci. 6:385-400.

. 1924. Anomalies in the distribution of fossil gulls.

Condor 24:173-174.

. 1925. Avifauna of the McKittrick Pleistocene. Univ.

Cal. Publ. Dept. Geol. Sci. 15(9):307-326.

. 1930. Further bird remains from the Upper San

Pedro Pleistocene. Condor 32: 1 1 6-1 1 8.

. 1931. Pleistocene birds from the Carpinteria as-
phalt of California. Univ. Cal. Publ. Bull. Dept. Geol.
Sci. 20:361-374.

. 1935. A second avifauna from the McKittrick

Pleistocene. Condor 37:72-79.

. 1944. A Pliocene flamingo from Mexico. Wilson

Bull. 56:77-82.

Miller, L.H., and I. DeMay. 1 942. Fossil birds of California.
Univ. Cal. Publ. Dept. Zool. 47(4):47- 1 42.

Sama-Wojcicki, A. 1980. Chemical analyses, correlations,
and ages of late Cenozoic tephra units of east-central
and southern California. U.S. Geological Survey Open
File Report 80:1-231.

Shufeldt, R.W. 1 892. A study of the fossil avifauna of the
Equus beds of the Oregon desert. Jour. Acad. Nat. Sci.
Phil. 9:389-425.

. 1913. A review of the fossils of the desert region

of Oregon, with a description of additional material col-

12 Contributions in Science, Number 362

Jefferson: Lake Manix Avifauna

lected there. American Mus. Nat. Hist. Bull. 32:1 23—
178.

Snyder, C.T., G. Hardman, and F.F. Zdenek. 1964. Pleis-
tocene lakes in the Great Basin. U.S. Geol. Survey, Misc.
Geol. Invest. Map 1-416.

Stock, C. 1953. Rancho La Brea. A record of Pleistocene
life in California, 5th ed. Los Angeles Co. Mus. Sci. Ser.
no. Paleon. 1 1: 1-79.

Wetmore, A. 1956. A check-list of the fossil and prehistoric
birds of North America and the West Indies. Smithson-
ian Misc. Coll. 131(5): 1-105.

Winters, H.H. 1954. The Pleistocene fauna of the Manix
Beds in the Mojave Desert, California. Dept. Geol. un-
publ. masters thesis, Calif. Inst. Technology, i + 58.

Accepted 6 December 1984.

Contributions in Science, Number 362

Jefferson: Lake Manix Avifauna 13

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FOSSIL PONTOPORIID DOLPHINS (MAMMALIA: CETACEA)
FROM THE PACIFIC COAST OF NORTH AMERICA

Lawrence G. Barnes1

ABSTRACT. Some remarkably long-snouted fossil marine odon-
tocete cetaceans of late Tertiary age from the eastern margin of the
North Pacific Ocean comprise an extinct group classified as the
subfamily Parapontoporiinae of the family Pontoporiidae. These
fossil dolphins belong in the genus Parapontoporia Barnes, 1984,
which now includes three known species: latest Miocene Parapon-
toporia pacifica Barnes, 1984, from Baja California, Mexico; latest
Miocene P. wilsoni, new species, from central California, U.S.A.;
and Late Pliocene P. sternbergi (Gregory and Kellogg, 1927), from
southern California. Analysis of the morphology of these animals
indicates that they are closely related to species that have been pre-
viously classified in two separate living families of dolphins: the
marine Pontoporiidae of the southwest Atlantic Ocean, and the
freshwater Lipotidae of China.

The living pontoporiid, the La Plata dolphin or fraciscana, Pon-
toporia blainvillei (Gervais and d’Orbigny, 1844), lives in shallow
water off the Atlantic coast of South America. It is apparently most
closely related to two fossil Pliocene marine species, Pontistes rec-
tifrons (Bravard, 1884) from Argentina and Pliopontos littoralis de
Muizon, 1 983, from Peru, and together these comprise the nominate
subfamily Pontoporiinae of the Pontoporiidae. The name-bearer of
the Lipotidae, the beiji or white flag dolphin, Lipotes vexillifer Miller,
1918, inhabits Tungting Lake and the Yangtze River (Chang Jiang)
in China. This species was traditionally classified in the family In-
iidae with the living Amazon dolphin or bouto, Inia geoffrensis (de
Blainville, 1817), and some fossil taxa, until 1979 when it was placed
in the monotypic new family Lipotidae.

The fossil species in the subfamily Parapontoporiinae are inter-
mediate morphologically and zoogeographically, however, between
L. vexillifer and the species in the subfamily Pontoporiinae, and
separate family status for L. vexillifer is therefore unwarranted. Based
on the new information from the fossil record I recognize, at a new
rank and in a new context, a third subfamily in the family Ponto-
poriidae, the Lipotinae.

INTRODUCTION

Four living genera of odontocetes in the superfamily Plata-
nistoidea contain long-snouted species that are sometimes
(although not in every case correctly) referred to as “river
dolphins.” Pontoporia blainvillei (Gervais and d’Orbigny,
1844), the franciscana or La Plata dolphin, is a near-shore
marine species living in the Atlantic Ocean off the coasts of

Contributions in Science, Number 363, pp. 1-34
Natural History Museum of Los Angeles County, 1985

Brazil, Uruguay, and Argentina. Inia geoffrensis (de Blain-
ville, 1817), the bouto or Amazon dolphin, is a freshwater
species living in the Amazon, Orinoco, and Madeira River
systems of South America. Lipotes vexillifer Miller, 1918,
the Beiji or white flag dolphin, is a freshwater dolphin in-
habiting the Yangtze (Chang Jiang) River system (including
Tungting Lake) and the Quiantang River system in China.
Platanista spp., the susus or Ganges and Indus dolphins of
India, Pakistan, Bangladesh, and Nepal are freshwater species.

Each of these four Recent genera has been designated as
the type of a separate platanistoid family, although in most
published works they have been classified in various subfam-
ilies within the family Platanistidae (e.g.. Flower, 1869;
Simpson, 1945; Fraser and Purves, 1960). Pontoporia Gray,
1846, and Platanista Wagler, 1830, have usually been clas-
sified in their nominate subfamilies, but following Miller
(1918, 1923), most zoologists have joined Lipotes Miller,
1918, and Inia d’Orbigny, 1834, in the family Iniidae or the
subfamily Iniinae, depending on whether the Platanistoidea
or Platanistidae was used as the next higher category (see
Simpson, 1945; Brownell and Herald, 1972). Some authors
have also classified Pontoporia within the Delphinidae (Gill,
1 872; True, 1908; Miller, 1 923; Kellogg, 1928), but they have
been in the minority. Zhou, Qian, and Li (1979) recognized
substantial differences between Inia and Lipotes and pro-
posed a separate new family, the Lipotidae. Several fossil
species, virtually all of them marine, have been reported to
be relatives of various of these, and the systematics of the
fossil and Recent Platanistoidea have had a convoluted and
confusing history (Kellogg, 1928).

In recent years a larger number of cetologists and pale-
ontologists have realized that the platanistoid dolphins are
morphologically diverse and represent more than one family.
For example, Kasuya (1973) recognized three extant families
(Platanistidae, Iniidae, and Pontoporiidae) and Zhou (1982)
recognized four, including the family Lipotidae. Both of these

1. Vertebrate Paleontology Section, Natural History Museum of
Los Angeles County, 900 Exposition Blvd., Los Angeles, California
90007.

ISSN 0459-8113

authors classified all of these families in the superfamily Plat-
anistoidea, so that the hierarchical relationships among the
groups remained virtually unchanged.

The purpose of this paper is to describe, diagnose, and
analyze fossils of extremely long-snouted marine dolphins in
the genus Parapontoporia Barnes, 1984, cetaceans that are
morphologically similar to both Pontoporia and Lipotes. That
Parapontoporia is morphologically intermediate between the
Recent genera Pontoporia and Lipotes is important to sys-
tematics because, as mentioned above, Pontoporia and Plat-
anista have usually been considered to be more remotely
related to Inia and Lipotes than the latter were to one another.

The fossils that I describe in this study include those of
Parapontoporia pacifica Barnes, 1984; P. sternbergi (Gregory
and Kellogg, 1927); and P. wilsoni, new species, that were
collected from latest Miocene and Late Pliocene sedimentary
rock units bordering the eastern North Pacific Ocean in Cal-
ifornia and Baja California. The type species of Paraponto-
poria, P. pacifica, was based on a fossil from latest Miocene
rocks, approximately 6 to 8 million years old, on Isla Cedros,
Baja California, Mexico. I have previously demonstrated
(Barnes, 1984) that P. pacifica is congeneric with Stenodel-
phis sternbergi Gregory and Kellogg, 1927, a species that is
known from the Late Pliocene age San Diego Formation in
California and one that had needed a new generic allocation
for many years (see Barnes, 1973a, 1977). The geographic
distribution of these fossils has important biogeographic im-
plications because they are roughly equidistant between the
areas occupied by Lipotes vexillifer and Pontoporia blainvil-
lei. These specimens and taxa were included in a Ph.D. dis-
sertation that I submitted to the University of California at
Berkeley (Barnes, 1972), and some were subsequently men-
tioned in four following publications (Barnes, 1973a, 1977,
1983, 1984). In a subsequent study, Robert L. Brownell, Jr.,
Edward Mitchell, and I plan to review all fossil and Recent
Pontoporiidae of the world and their interrelationships.

MATERIALS AND METHODS

Fossil specimens described in this study are conserved in
scientific institutions in the United States as indicated by the
following acronyms:

AMNH American Museum of Natural History, New Y ork,
New York.

LACM Natural History Museum of Los Angeles County,
Los Angeles, California.

SDSNH San Diego Society of Natural History, Natural
History Museum, San Diego, California.

UCMP University of California Museum of Paleontology,

Berkeley, California.

UCR University of California at Riverside, Department
of Geological Sciences, Riverside, California.

Precise locality descriptions are not given for some of the
specimens. Such information is available to qualified in-
vestigators upon request to the appropriate institution.

Comparative specimens of the modem species, Pontoporia
blainvillei, Lipotes vexillifer, Inia geojfrensis, and Platanista

gangetica, were studied and used in formulating the descrip-
tions and diagnoses, which are based on cranial characters.
Each of the three fossil species of Parapontoporia is known
by incomplete skulls that do not exhibit entirely overlapping
morphology, but enough is known of each species to differ-
entiate all three. The descriptions presented here do not du-
plicate those already published, but are written in a manner
to avoid repetition. Measurements of the skulls were taken
following the standardized methods outlined by Perrin (1975).
Anatomical terminology for basicranial structures and the
middle ear air sinus system follows Fraser and Purves ( 1 960).
Most other osteological terms follow Keman (1918), Kellogg
(1927), and Bames (1978, 1984). All the rendered antomical
line drawings with line shading (Figs, la, 2a, 3a, 4, 10a, b,
11, 13, 15) were done by J. Patricia Lufkin using orthographic
projection. My restorations of Parapontoporia wilsoni were
derived from these. Specimens were coated for photography
with a sublimate of ammonium chloride. My restorations of
P. sternbergi are based on all available specimens from the
San Diego Formation and on the type specimens of the other
two species.

Anatomical structures in the illustrations are labeled ac-
cording to the following abbreviations:

aon— antorbital notch
Bs— basisphenoid bone
ch— cranial hiatus
fc— carotid foramen
fh — hypoglossal foramen
fio— infraorbital foramen
fmx — maxillary foramen
fo— foramen ovale

fp— falcate process of the basi occipital

fpal — palatine foramen

fpmx— premaxillary foramen

Fr— frontal bone

gf— glenoid fossa

jn— jugular notch

Ju —jugal bone

La — lacrimal bone

me— maxillary crest

me— maxillary eminence

Met— mesethmoid bone

mf— mental foramen

mrg— mesorostral gutter

ms— fossa for middle sinus

Mx— maxillary bone

n — naris

Na— nasal bone

Oc— occipital bone

occ— occipital condyle

Pa— parietal bone

Pal — palatine bone

pop— paroccipital process

Pmx— premaxillary bone

Pt(ll) — • pterygoid, lateral lamina

Pt(ml)— pterygoid, medial lamina

pts— fossa for pterygoid sinus

2 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

s— mandibular symphysis
Sq— squamosal bone
sqf— squamosal fossa
Vo— vomer bone

Some family group names have been used at various ranks.
In cases where I recognize a revised rank, the original author
is included in parentheses followed by the author of the
emended rank that I recognize. All the variations and syn-
onyms of the family- and genus-group names are listed in
the synonymies.

SYSTEMATICS

Class Mammalia Linnaeus, 1758
Order Cetacea Brisson, 1762
Suborder Odontoceti Flower, 1867

Superfamily Platanistoidea (Gray, 1863)
Simpson, 1945

Family Pontoporiidae (Gill, 1871)

Kasuya, 1973

Pontoporiinae Gill, 1871:124 (March, 1871), as a subfamily
of Platanistidae; 1872:15, as a subfamily of Delphinidae.
Pontoporiadae Gray, 1871:95, incorrectly formed name, as
a monotypic family for Pontoporia Gray, 1846.
Stenodelphinae True, 1908:391, as a subfamily of Delphin-
idae to include Stenodelphis d'Orbigny and Gervais, 1 847,
a junior synonym of Pontoporia.

Stenodelphininae. Miller, 1923:40, unjustified emendation
of Stenodelphinae.

Pontoporiidae. Kasuya, 1973:28, 61, as a family of Plata-
nistoidea.

Lipotidae Zhou, Qian, and Li, 1 979:72, as a monotypic fam-
ily for Lipotes.

DIAGNOSIS OF FAMILY. A family of Platanistoidea
that differs from Platanistidae and Iniidae by having skulls
with the following unique combination of characters: occip-
ital shield roughly square in posterior view with prominent
dorsolateral comers and not narrow dorsaily, facial portions
of maxillae relatively flat, not steeply inclined posteriorly or
laterally, rostral portions of premaxilla and maxilla separated
by longitudinal lateral groove, rostrum constricted trans-
versely posterior to end of tooth row, posterior part of al-
veolar row located at lateral edge of palate and curving up-
ward on side of rostrum at posterior extremity, palate nearly
flat posterior to ends of alveolar rows, maxilla bearing a low,
non-pneumaticized crest over orbit which is oriented in an
anterolateral to posteromedial direction and located medial
to margin of maxilla rather than at the margin, fossa for
pterygoid sinus having a branch extending dorsaily adjacent
to anterior wall of each naris, nares of small diameter and
curving around anterior wall of braincase rather than being
nearly vertical, foramen ovale small, round, distinct from
cranial hiatus, located on basisphenoid bone on ventrolateral

wall of cranium and confluent with deep sulcus marking
course of mandibular division of trigeminal nerve, zygomatic
process of squamosal elongate, tapered, inclined anteriorly
with large, transversely oriented postglenoid process, and
prominent, anteroposteriorly oriented fossa curving around
medial and posterior side of glenoid fossa for middle sinus
of air sinus system, paroccipital process with fossa on side
facing cranial hiatus that held posterior sinus of air sinus
system, carotid foramen in basioccipital vestigial, periotic
and bulla not firmly attached to braincase by posterior pro-
cesses and accessory ossicles, teeth comparatively homodont
with crowns having a lingual protuberance and roots having
a swelling below the enamel line.

INCLUDED SUBFAMILIES. Parapontoporiinae Barnes,
1984; Pontoporiinae (Gill, 1871) Kasuya, 1973; and Lipo-
tinae (Zhou, Qian, and Li, 1 979), new rank and new context.

Subfamily Parapontoporiinae Barnes, 1984

Parapontoporiinae Barnes, 1984:6.

EMENDED DIAGNOSIS OF SUBFAMILY. A subfam-
ily of Pontoporiidae differing from Pontoporiinae and resem-
bling Lipotinae by having skull with cranial vertex asym-
metrical and offset to left side, vomer exposed between
maxillae on palate, spiracular plate on premaxillary surfaces
flat, not elevated and convex, posterior terminations of pre-
maxillae not widely separated from anterolateral corners of
nasals, squamosal fossa between zygomatic process and
braincase deep; resembling Pontoporiinae and differing from
Lipotinae by having skull with extremely long, slender ros-
trum and mandible, very deep lateral groove between rostral
parts of maxilla and premaxilla, lateral lamina of pterygoid
joined with posterior plates of maxilla and palatine to form
extensive bony wall within orbit (but not connecting poste-
riorly with basisphenoid as in Pontoporia blainvillei ), tooth
crowns small, sharply pointed, with basal lingual bulge, com-
pressed anteroposteriorly and covered with smooth enamel,
tooth roots flattened labio-lingually with encircling swelling
at gum line; and differing from both Pontoporiinae and Li-
potinae by having extreme polydonty, bearing 80 to 82 teeth
in each side of each jaw (in contrast to 48 to 6 1 in Pontoporia
blainvillei and 32 to 36 in Lipotes vexillifer).

INCLUDED GENERA. Parapontoporia Barnes, 1984
only.

Parapontoporia Bames, 1984

Stenodelphis (part). Gregory and Kellogg, 1927.

“ Stenodelphis Bames, 1973a:37-39; 1977:333-334; Bames,
Howard, Hutchison, and Welton, 1981:56, 57, 61, 64; De-
mere, 1981:24-25; de Muizon, 1983:1 103.
Stenodelphininae, genus (and species) new. Bames, 1977:
331.

Parapontoporia Bames, 1984:6.

DIAGNOSIS OF GENUS. Because the subfamily Para-
pontoporiinae is at present monotypic, the diagnosis for it
and the genus Parapontoporia are identical.

Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins 3

TYPE SPECIES. Parapontoporia pacifica Barnes, 1984;
type by original designation.

INCLUDED SPECIES. Parapontoporia sternbergi (Greg-
ory and Kellogg, 1927); Parapontoporia pacifica Barnes, 1984;
and Parapontoporia wilsoni, new species.

Parapontoporia pacifica Barnes, 1984

Figures 1-5, 20a, d

Stenodelphininae, genus and species new (part). Barnes, 1977:

331.

Parapontoporia pacifica Barnes, 1984:7.

EMENDED DIAGNOSIS OF SPECIES. A species of
Parapontoporia differing from P. wilsoni, new species, and
P. sternbergi by having skull with dorsal premaxillary sur-
faces at proximal end of rostrum flat-lying, not tilting me-
dially to form a basin, and by having more elongate fossa
for pterygoid sinus in palatine; and differing from P. stern-
bergi by lacking well-defined grooves on the lateral surfaces
of dentaries, instead, nutrient foramina on lateral surface of
dentary open into only a shallow, elongate depression.

HOLOTYPE. UCR 2 1 244, skull with teeth, lacking the
braincase and part of the facial region, mandible lacking right
condyle and the post-alveolar section of the left dentary, a
probable fifth cervical vertebra, and a first left rib; collected
by David P. Whistler, 9-1 1 August 1965.

TYPE LOCALITY. UCR RV-6505 in weathered bad-
lands exposures southeast of the village of Cedros on Isla
Cedros, Baja California, Mexico.

FORMATION AND AGE. From the lower member of
the Almejas Formation (Kilmer, 1979), latest Miocene, cor-
related indirectly with the “Jacalitos” provisional mega-in-
vertebrate stage and the Hemphillian North American land
mammal age, and approximately 6 to 8 million years old.
The age of the lower member of the Almejas Formation has
been discussed by Barnes (1973b, 1977, 1984) and Repen-
ning and Tedford (1977). The Almejas Formation is ap-
proximately 800 feet thick and directly overlies the Middle
Miocene age Tortuga Formation (Kilmer, 1977, 1979). Fossil
invertebrate assemblages from the upper member of the Al-
mejas Formation are similar to those from the Late Pliocene
San Diego Formation and are therefore correlative with the
“San Joaquin” provisional mega-invertebrate stage and the
Blancan North American land mammal age. The vertebrate
fossils in the lower member of the Almejas Formation on
Isla Cedros are stratigraphically below the invertebrate fos-
sils, in the basal approximately 1 30 feet of the formation.

The holotype of Parapontoporia pacifica was found five
feet above the base of the Almejas Formation. Howard (1971)
reported seven species ofbirds. Repenning and Tedford (1977)
described walruses and a fur seal, and Barnes (1973b, 1977,
1984) reported 1 1 species of cetaceans from the lower mem-
ber of the formation, mostly from horizons above that which
produced P. pacifica.

The relatively diverse aggregate assemblage of published
fossil vertebrate species from the lower member of the Al-
mejas Formation warrants being listed here:

Aves

Pujfinus tedfordi Howard, 1971
Puffin us sp.

Morus sp.

t Mega p aloe l od us opsigonus Brodkorb, 1961
Cerorhinca minor Howard, 1971
lEndomychura sp.

Mancalla cedrosensis Howard, 1971
Mammalia

Dusignathus santacruzensis Kellogg, 1 927
Aivukus cedrosensis Repenning and Tedford, 1977
Thalassoleon mexicanus Repenning and Tedford, 1977
cf. Plesiocetus sp.

Balaenoptera sp.

Parapontoporia pacifica Barnes, 1984
Denebola brachvcephala Barnes, 1984
Piscolithax tedfordi Barnes, 1984
Piscolithax boreios Barnes, 1984
Phocoenidae, gen. and sp. undetermined
Albireo whist/eri Barnes, 1984
Delphinoidea, gen. and sp. undetermined
Praekogia cedrosensis Barnes, 1973
Physeterinae, gen. and sp. undetermined

DESCRIPTION. Skull. The skull (Figs. 1-3) of the ho-
lotype and only known specimen of Parapontoporia pacifica
was described by Barnes (1984) and compared with several
specimens of P. sternbergi from the younger. Late Pliocene
age San Diego Formation. The relatively complete referred
specimens of P. sternbergi described in the following text
confirm that the two species are congeneric. The description
of the skull of P. pacifica need not be repeated here, but I
will summarize some of the main differences between it and
the other species of Parapontoporia. The holotype of P. pa-
cifica does not include the posterior part of the braincase,
but it has the most complete rostrum of any known specimen
of the genus. No available specimen of P. sternbergi has the
tip of the rostrum preserved.

Some of the lateral lamina of the pterygoid of P. pacifica
is preserved, and in company with a small posterior extension
of the palatine and a larger projection of the maxilla, produces
a thin wall of bone that extends posteriorly within the orbit
(Fig. 2). This wall of bone spreads dorsally and partly ob-
scures the foramina and sinuses on the medial wall of the
orbit, but it does not appear to have reached posteriorly as
far as the basisphenoid as it does in Pontoporia blainvillei.
This is the only specimen of Parapontoporia in which any
part of the lateral lamina of the pterygoid is preserved.

The groove on the lateral side of the rostrum that separates
the premaxilla from the maxilla is not as deep as in P. stern-
bergi. The groove becomes increasingly wider and shallower
distally, and at the anterior end of the rostrum the maxilla
is fused to the premaxilla with no visible suture. Both P.
sternbergi and P. wilsoni, new species, have a medial basin
formed on the dorsal surface of the proximal part of the
rostrum just anterior to the level of the antorbital notches.
In P. pacifica, there is no such basin because the premaxillae
are nearly flat-lying in that area, but both their medial and

4 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

CD

JQ

Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins 5

Figure 1. Parapontoporia pacifica Barnes, 1984, holotype, UCR 21244, partial skull, dorsal view: a, drawing of original specimen; b, restoration with structures labeled.
Abbreviations used in this and the following illustrations are explained in the section titled Materials and Methods.

6 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

Figure 2. Parapontoporia pacifica Barnes, 1984, holotype, UCR 21244, partial skull, right lateral view: a, drawing of original specimen; b, restoration with structures labeled.

Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins 7

Figure 3. Parapontoporia pacifica Barnes, 1984, holotype, UCR 21244, partial skull, ventral view: a, drawing of original specimen; b, restoration with structures labeled.

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lateral margins are very slightly upturned (Figs. 20a-c). At
this location there is no narrowing of the premaxillae as in
P. wilsoni, new species.

The fossae in the palatine bones that held the pterygoid
air sinuses are as long anteroposteriorly as in P. sternbergi,
and nearly twice the length of those in P. wilsoni, new species.
The rostrum had approximately 80 teeth on each side.

Mandible. The mandible with the holotype of P. pacifica
(Fig. 4) is the most complete known mandible of any species
of Parapontoporia. The right and left dentaries each bore
alveoli for approximately 82 teeth. The specimen shares
characters with the holotype mandible (AMNH 2 1 905) of P.
sternbergi and is similar in general size and proportions. It
has a firmly fused mandibular symphysis marked by a medial
groove dorsally and ventrally. The mandible of P. pacifica
differs from that of P. sternbergi by lacking a deep lateral
groove (such as is present in Recent Pontoporia blainvillei ),
but has instead only an elongate shallow depression that
extends the length of the symphyseal part of each dentary
slightly below mid-height on the lateral side (compare Figs.
20d and e).

All bone surface of the mandible is dense and in its tooth-
bearing parts the mandible is marked by faint linear striae.
Between the alveolar row and the dorsal edge of the lateral
groove, the surface of the dentary is bowed outward in cross
section. The dorsal surface of each dentary between the mid-
line and the alveolar row is arched transversely.

The ascending ramus has a thin, arched coronoid crest
about mid-length, and this culminates posteriorly in an up-
turned coronoid process (Fig. 4b). Neither Lipotes vexillifer
nor Pontoporia blainvillei has a coronoid crest and in both
species the coronoid process projects more posteriorly than
dorsally.

Teeth. The teeth of Parapontoporia pacifica are like those
of Pontoporia blainvillei (Figs. 5a-d), but have relatively higher
crowns. The smooth enamel crowns curve lingually, have a
basal lingual bulge, and are compressed anteroposteriorly.
The roots have a swelling encircling them at the gum line
and are compressed labio-lingually. The anterior teeth have
relatively slender, erect crowns, but progressing posteriorly
the crowns are shorter and bend more medially. Because they
are more bulbous, the crowns of the posterior teeth are not
as compressed anteroposteriorly as are the ones of the more
slender anterior teeth.

Cervical vertebra. One isolated vertebra (Fig. 5e), probably
a fifth cervical vertebra, was associated with the holotype
skull and mandible. This vertebra is greatly compressed
anteroposteriorly, has a centrum that is nearly square in out-
line, a slender neural arch, small diapophyses and parapophy-
ses, and a large transverse foramen that probably did not
have a complete bony arch on its lateral side.

This vertebra resembles the fifth cervical of Lipotes vex-
illifer (Miller , 1 9 1 8:pl. 12, the vertebra that is placed at lower
left in the illustration) in having a low, broad neural canal,
dorsoventrally expanded transverse foramen, and ventrally
directed parapophyses. The resemblance of this vertebra to
the fifth cervical of Pontoporia blainvillei is not so striking,
because in that species (de Carvalho, 1961 :fig. 1 le) the bone

8 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

Figure 5. Parapontoporia pacifica Barnes, 1984, holotype, UCR 21244, right mandibular tooth: a, lingual view; b, anterior view; right
maxillary tooth: e, lingual view; d, anterior view; e, cervical vertebra, anterior view; f, anterior right rib, anterior view. Scale line is for Figs,
a-d only. Figs, e, f natural size.

has relatively larger diapophyses and parapophyses, a higher
neural canal, a dorsoventrally compressed transverse fora-
men, a broader centrum, and no ventral extension of the
parapophyses.

Rib. The proximal part of one rib (Fig. 50, apparently a
first left rib, belongs with the holotype. This rib has a large
capitulum, a flat tuberculum, a short neck, a wide but antero-
posteriorly compressed body, and is strongly curved at its
proximal end.

Parapontoporia sternbergi
(Gregory and Kellogg, 1927)

Figures 6-10, 17b, 20c, e, 21b

Stenodelphis sternbergi Gregory and Kellogg, 1927:1.

“ Stenodelphis ” sternbergi Gregory and Kellogg, 1927. Bames,
197 3a:37— 38; 1977:333-334; Bames, Howard, Hutchison,
and Welton, 1981:56, 57, 61, 64; Demere, 1981:24-25; de
Muizon, 1983:1 103.

Parapontoporia sternbergi (Gregory and Kellogg, 1927).
Bames, 1984:6.

EMENDED DIAGNOSIS OF SPECIES. A species of
Parapontoporia differing from P. wilsoni, new species, by
having skull with facial region wider than long, antorbital
processes larger, cranial vertex (comprised of frontals and
nasals) higher and more compressed transversely, occipital

condyles projecting less prominently posteriorly from occip-
ital shield, premaxillary surfaces forming less of a basin at
proximal end of rostrum, nares passing more vertically
through the skull, temporal fossa compressed anteroposte-
riorly with its height and length approximately equal, more
elongate fossa for pterygoid sinus in palatine, zygomatic pro-
cess of squamosal more inclined anteriorly, squamosal be-
tween postglenoid process and paroccipital process highly
compressed anteroposteriorly; and differing from P. pacifica
by having basin formed on premaxillary surfaces on proximal
part of rostrum, shorter fossa for pterygoid sinus in palatine,
and deeper groove on rostrum between maxilla and pre-
maxilla, and by having mandible with deep groove on lateral
side of each dentary.

HOLOTYPE. AMNH 2 1 905, a section of mandible in the
symphyseal area with 24 teeth which have badly eroded
crowns, collected by John Reiland. The specimen is not a
rostral fragment as was stated by Gregory and Kellogg ( 1 927).

TYPE LOCALITY. India Street at West Walnut, San Die-
go, San Diego County, California.

REFERRED SPECIMENS FROM THE SAN DIEGO
FORMATION. Nearly complete skulls, LACM 6238,
SDSNH 22633, 23084; rostra lacking braincases, LACM
26605, SDSNH 25022; cranial vertices, LACM 58902, UCMP
129662, UCMP 129663; cranial vertex associated with ros-
tral fragments, LACM 30464; right squamosal, LACM 26597;
partial left maxilla, SDSNH 24794; cranial fragments, LACM

Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins 9

30465; periotics, LACM 58901, LACM 103975, SDSNH
20941, 23058, 23630, 23631, 24734, 25049, UCMP 88581,
UCMP 88589, UCMP 129660, UCMP 129661; rostral frag-
ments associated with mandible fragments and teeth, UCMP
88590; premaxillary fragment with teeth, UCMP 88588; ros-
tral fragments, LACM 30432, LACM 30433, LACM 31065;
nearly complete mandible, SDSNH 18671; and mandible
fragments, LACM 26594, LACM 30463, LACM 30468,
LACM 30471 (with teeth), LACM 30472, LACM 30475,
LACM 30476, LACM 104004, UCMP 88587.

FORMATION AND AGE. Lower member of the San
Diego Formation, Late Pliocene, correlated with the “San
Joaquin” provisional mega-invertebrate stage of Addicott
(1972) and the Blancan North American land mammal age,
and therefore approximately 2 to 4 million years old. The
fossil fauna and age of this rock unit have been discussed by
Howard (1949), Hertlein and Grant (1954), Barnes (1973a,
1977:332-334), Repenning and Tedford (1977:69-70, fig. 6),
and Demere (1982, 1983).

DESCRIPTION. Skull. The rostrum, mandible, and teeth
are the only anatomical parts shared in common between
known specimens of P. pacifica and P. sternbergi, and the
braincase and proximal part of the rostrum are the only parts
shared in common between specimens of the latter species
and P. wilsoni, new species. To avoid repetition, I shall em-
phasize in the following text those characters whereby P.
sternbergi differs from P. wilsoni, new species, and P. pacifica,
and those parts of the skull that are not known for P. pacifica.

The skull of P. sternbergi (Figs. 6-8) has a braincase that
is somewhat square or boxy in its proportions and it has an
extremely long and narrow rostrum. Many of the characters
that separate P. sternbergi from P. wilsoni, new species, are
related to the derived state of foreshortening or anteropos-
terior compression of the braincase of P. sternbergi (Fig. 21).
This compression is the phenomenon termed telescoping by
Miller (1923), whereby the crania of Cetacea have become
modified from the condition in generalized mammals so that
the bones extend anteroposteriorly past one another. In
odontocetes, the predominance of posterior movement of
the rostral bones toward the occipital region is typical, and
in the more derived states, involved an anteroposterior
compression of the braincase as well.

The braincase proportions of P. sternbergi are somewhat
like those of Recent Lipotes vexillifer (Figs. 1 7b, c), but are
different in details, both primitive and derived. The braincase
is deep dorsoventraily and has a nearly vertical occipital
shield and a wide and fairly flat facial surface that narrows
abruptly at the juncture with the rostrum.

On the posterior part of the rostrum, the dorsal surfaces
of the premaxillae slope medially toward the mesorostral
gutter to form a central basin. At its deepest point, just an-
terior to the antorbital notches, this basin is approximately
5 mm lower than the adjacent maxillary surfaces (Fig. 20c).
Progressing anteriorly from the basin the premaxillae grad-
ually become less tilted. There is a triangular shaped rough
area on each premaxilla anterior to the nares. In delphinoids
this is the site of attachment of the nasal plug muscle (Law-
rence and Schevill, 1956), and a similar situation undoubt-

edly existed in P. sternbergi. The narrow anterior end of this
rough area extends into the deepest part of the rostral depres-
sion (Fig. 6).

For most of their rostral length the premaxillae are slender,
very dense and are separated from the maxillae by a deep
longitudinal groove on each side (Fig. 7). There are no ob-
vious maxilla-premaxilla sutures within these grooves. The
premaxillae form most of the dorsal surface of the rostrum
and roof over the mesorostral gutter, touching (but not fused)
for much of their length at the midline. They extend to the
extreme anterior tip of the rostrum, and are separated me-
dially there for a short distance to expose the mesorostral
gutter. Near the rostral basin the premaxillae become in-
creasingly wider, the grooves between them and the maxillae
disappear, and the premaxillae diverge to expose the me-
sorostral gutter.

At a point approximately 5 mm posterior to the antorbital
notches, each premaxilla is perforated by a premaxillary fo-
ramen which, as in a common odontocete pattern, is con-
nected to three sulci; anteromedial, posteromedial, and pos-
terolateral. The anteromedial sulcus is partly roofed over by
bone and defines the lateral margin of the narrow anterior
extension of the rugose area of attachment of the nasal plug
muscle (mentioned above). The posterolateral sulcus is deep-
ly incised into the premaxilla. It diverges sharply away from
the midline of the skull as it courses toward the lateral margin
of the premaxilla, then continues posteriorly in a shallow
groove on the maxilla following the edge of the premaxilla.
The posteromedial sulcus branches off the posterolateral sul-
cus just posterior to the premaxillary foramen, but it is very
shallow and extends only approximately 15 mm.

Beginning immediately anterior to the nares, the premax-
illae become slightly elevated above the surrounding max-
illae, and this condition persists to the posterior termination
of each premaxilla. These elevated areas around the nares
were smooth surfaced in life, but on either side of the avail-
able skulls the bone surface is now partly eroded away. When
the animal was alive, these slightly elevated premaxillary
surfaces, the spiracular plates, supported the premaxillary
sacs (which are diverticula of the nasal passages, see Mead,
1975). These plates are asymmetrical. The one on the right
side is wider and more elevated on its lateral edge than the
one on the left. The left plate tilts more toward its lateral
edge, and extends farther posteriorly. Each premaxilla ter-
minates about 5 mm from the corresponding nasal. Extend-
ing posteriorly from the posterior end of each premaxilla
there is a rough area on the maxilla indicating that at some
previous point in the evolutionary history of this lineage of
dolphin the premaxilla had extended farther posteriorly ad-
jacent to the nasal as in, for example, the squalodonts.

The mesethmoid septum between the nares is canted to
the left as it rises from the skull, and the left naris is slightly
larger than the right. The basic construction of the bones
surrounding the nares is very similar to that in Lipotes vex-
illifer, and is different from that in Pliopontos littoralis and
Pontoporia blainvillei, in which the spiracular plates are more
elevated and the bones around the nares are symmetrical
(Fig. 17).

10 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins 11

Figure 6. Paraponloporia sternbergi (Gregory and Kellogg, 1927), referred specimen, LACM 6238, skull, dorsal view: a, photograph of original specimen; b, restoration based on
all known specimens of the species with extremity of rostrum based on P. pacifica.

12 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

Figure 7. Parapontoporia sternbergi (Gregory and Kellogg, 1927), referred specimen, LACM 6238, skull, left lateral view: a, photograph of original specimen; b, restoration based
on all known specimens of the species with extremity of rostrum based on P. pacifica.

Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins 13

Figure 8. Parapontoporia sternbergi (Gregory and Kellogg, 1927), referred specimen, LACM 6238, skull, ventral view: a, photograph of original specimen; b, restoration based
on all known specimens of the species with extremity of rostrum based on P. pacifca.

Table 1. Measurements (in mm) of skulls of Parapontoporia. Parentheses indicate estimated measurements. Method follows Perrin (1975).

P. pacifica

P. wilsoni

P. sternbergi

LJCR

21244

UCMP

83790

LACM

6238

LACM

26605

SDSNH

22633

SDSNH

23084

Condylobasal length

—

—

(596 + )

—

(605 + )

—

Length of rostrum

477

-

(467 + )

(341+)

(475 + )

-

Width of rostrum at base

(80)

71

69

(64)

(64 + )

(61+)

Width of rostrum at midlength

17

—

—

—

—

—

Width of premaxillae at midlength of rostrum

12

-

-

—

—

—

Greatest preorbital width

(126)

104

108

—

—

—

Greatest postorbital width

-

(124)

-

—

120

(118)

Least supraorbital width

(112)

113

—

—

—

—

Greatest width of external nares

—

33

31

—

26

26

Width across zygomatic processes of squamosals

-

138

141

—

(136)

—

Greatest width of premaxillae

-

58

58

—

50

(54)

Greatest parietal width within temporal fossae

-

106

109

—

98

—

Vertical external height of brain case

-

—

94

—

85

—

Internal length of brain case

-

—

94

—

—

—

Length of temporal fossa

-

(73)

-

—

(72)

—

Width of temporal fossa

-

(48)

—

-

41

_

Length of orbit

-

44

—

—

—

—

Length of antorbital process of lacrimal

22

22

-

-

—

—

Length of tooth row

414

—

(398+)

(282 + )

(430 + )

-

Like the premaxillae, the maxillae are for much of their
rostral length composed of dense bone on both their labial
and palatal surfaces. The rostrum is constricted transversely
at a point approximately 50 mm anterior to the antorbital
notches. Both Lipotes vexillifer and Pontoporia blainvillei
have a similar constriction, and it is formed in the following
manner. In this area the dense part of the maxilla which is
lateral to the tooth row departs from the edge of the pre-
maxilla and extends onto the ventrolateral edge of the ros-
trum. Posterior to this and anteromedial to the antorbital
notches, the porous part of the maxilla is elevated into an
eminence that swells dorsally and laterally. The tapered an-
terior end of this maxillary eminence extends anteriorly
forming the lateral edge of the rostrum and passes dorsal to
the posterior end of the dense section of the maxilla forming
the tooth-bearing part of the rostral margin. This constriction
of the margin of the rostrum is typical of all Pontoporiidae
(Fig. 17).

The maxillary eminence has pushed the margin of the
adjacent premaxilla medially and also constricts the anterior
opening of the antorbital notch (Fig. 6). Posteromedial to
each maxillary eminence is a single maxillary foramen which
is confluent with shallow sulci running posteriorly and an-
teriorly from it, parallel to the margin of the premaxilla.

All known skulls of P. sternbergi have suffered breakage
and/or abrasion of the supraorbital area so that details of the
antorbital process and maxillary crest are not known for the
species. From what is preserved, the species appears to be
similar in these structures to P. pacifca and P. wilsoni, new

species, but to have had a more prominent antorbital process.
There is an obliquely oriented maxillary crest over the orbit,
and the skull SDSNH 22633 has the anterior end of the
zygomatic process of the jugal located anteromedial to the
antorbital notch (Demere, 1981:fig. 9) as in the other two
species. The same skull, illustrated by Demere, has the best
preserved postorbital process of the frontal of any specimen
known of the genus. The process is short and broadly tri-
angular, proportionally shorter and smaller than in Ponto-
poria blainvillei, and not slender with a distal rugosity as in
Lipotes vexillifer. It apparently did not contact the tip of the
zygomatic process of the squamosal (Fig. 7).

The posterior end of each maxilla wraps around the pos-
terior side of each nasal and contacts the elevated frontal on
the cranial vertex. In so doing, the maxillae encroach so far
medially upon the frontals that just posterior to the cranial
vertex only a 2-mm-wide exposure of frontals separates the
right maxilla from the left. The cranial vertex is formed of
the frontals and nasals and is in the form of an anteropos-
teriorly elongate, asymmetrical, slightly twisted knob with a
cleft at the median suture on its anterior surface. Like the
narial region, it is offset to the left side. The median suture
between the right and left frontals on the cranial vertex is 1 3
mm to the left of the midline of the skull, as marked by the
septum within the braincase that separated the cerebral hemi-
spheres of the brain. The exposed frontals form the highest
point of the vertex just posterior to the nasal bones, and there
is a lower area between that point and the occipital crest.
This low area posterior to the vertex exists also in other

14 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

Pontoporiidae and in Iniidae, and is unlike the condition in
species of Delphinidae and Phocoenidae, in which the cranial
vertex increases in height continuously from the nares to the
occipital crest. Each nasal is thin and wraps around the steep-
ly inclined anterior side of its corresponding frontal. The
mesethmoid, forming the posterior walls of the nares, is in-
clined in the same plane as the nasals, from which it is sep-
arated by arcuate sutures.

The occipital shield is oriented almost vertically (Fig. 7),
much as in Lipotes vexillifer. As in both Pontoporia blainvillei
and Lipotes vexillifer, it is approximately square in posterior
view, and adjacent to the temporal fossae its dorsolateral
corners are prominent. As in Pontoporia blainvillei, the mid-
line of the occipital shield is marked by a prominent median
sulcus that extends from the apex of the foramen magnum
dorsally to the occipital crest, and is flanked by a large, convex
area on each side corresponding to the cerebral hemispheres
of the brain. Each convex area is separated from the occipital
condyle below it by a prominent oblique sulcus that is not
present in Pontoporia blainvillei. The occipital condyles are
moderately convex and set off prominently from the occipital
shield. The dorsal margin of the foramen magnum forms a
triangular peak somewhat like that in Pontoporia blainvillei.
In Lipotes vexillifer the foramen magnum is more circular
in shape.

The temporal fossa is open posteriorly and curves around
the lateral side of the occipital shield. Dorsally, the facial
parts of the maxilla and frontal project approximately 8 mm
laterally over the posterior part of the temporal fossa. The
squamosal fossa, which floors the temporal fossa, forms a
deep recess between the cranium and the zygomatic process
of the squamosal. Anteriorly the squamosal fossa is floored
by only a thin shelf of bone spanning between the zygomatic
process and the cranium.

The paroccipital process is located relatively far anteriorly
on the braincase and therefore lies beneath the posterior end
of the temporal fossa. The paroccipital process terminates
ventrally in a flat, rugose surface and is separated medially
from the falcate process of the basioccipital by a narrow,
deep jugular notch. At the apex of this notch is a relatively
large hypoglossal foramen that is nearly 3 mm in diameter.

The zygomatic process of the squamosal is deep and long,
and its anterior end is upturned. There is a large, pointed,
ventrally directed postglenoid process. The glenoid fossa is
canted dorsomedially on the zygomatic process so that the
external surface forms a flange which projects ventrolaterally.
This ventrally projecting border is deepest anterior to the
center of the glenoid fossa and posterior to the point where
the zygomatic process turns upward. The glenoid fossa ex-
tends medially as a thin shelf which partly underlies a very
large fossa for the middle sinus of the middle ear air sinus
system. The fossa extends anteriorly along the medial side
of the zygomatic process much as in Lipotes vexillifer and
Pontoporia blainvillei. On its lateral surface the zygomatic
process of the squamosal is excavated posteriorly by a rugose
stemomastoid muscle fossa. The postglenoid process of the
squamosal is close to the paroccipital process and there is
much wrinkling and compression of the intervening bone,

including the muscle fossa, immediately dorsal to the exter-
nal auditory meatus. Such extreme anteroposterior compres-
sion does not exist in this part of the squamosal in P. wilsoni,
new species.

The cranial hiatus is the opening between the squamosal
and the basioccipital in which the periotic lay and through
which nerves and blood vessels passed, connecting the ear
to the endocranial cavity. This hiatus in P. sternbergi is large
(Fig. 8), but not relatively as large as in Pontoporia blainvillei.
There is no indication that there was a large falciform process
of the squamosal lateral to the hiatus such as is present in
species of Delphinidae. A small falciform process is present
in Pontoporia blainvillei. Large fossae in the bone around the
cranial hiatus of P. sternbergi indicate the locations in life of
extensive air sinuses. There was a large posterior sinus in the
anterior wall of the paroccipital process. A much larger one
occurs in the same location in Pontoporia blainvillei, and a
smaller one exists in Lipotes vexillifer. The peribullary sinus
in P. sternbergi extended laterally dorsal to the posterior
process of the periotic, as well as medially where it occupied
a large fossa in the lateral side of the falcate process of the
basioccipital. The extent of excavation of the falcate process
by this fossa is exceptional when compared to most other
species of odontocetes, especially Pontoporia blainvillei, in
which this process is very thick. At the front of this fossa,
the very small carotid foramen pierces the basioccipital ap-
proximately 5 mm posterior to the suture between the pter-
ygoid and the basioccipital. Lateral to this the basisphenoid
bridges between the squamosal and the pterygoid, and bears
the foramen ovale which is 4 mm in diameter and located
approximately 8 mm anterior to the anterior margin of the
cranial hiatus. A deep sulcus, marking the former course of
the mandibular division of the trigeminal nerve, leaves the
foramen ovale and extends posterodorsally across the basi-
sphenoid (Fig. 8).

The medial lamina of the pterygoid forms a crest that is
continuous with, but thicker than, the falcate process of the
basioccipital. The posterior end of the vomer covers the ven-
tral surface of the basisphenoid, spreads laterally to within
3 mm of the ventral edge of each pterygoid crest, and stops
at the basisphenoid-basioccipital suture. Posteriorly, each
falcate process of the basioccipital ends in a curved margin,
and between that and the paroccipital process of the exoc-
cipital the hypoglossal foramen lies at the apex of the jugular
notch. In Pontoporia blainvillei the same foramen lies outside
of, and posterior to the notch, rather than within it.

The anterolateral wall of the braincase is very well pre-
served only in LACM 6238. It is convex and remarkably
devoid of foramina, sinuses, and bony processes or crests.
The orbit is not as well preserved on any known specimens
of P. sternbergi as it is on the holotype of P. wilsoni , new
species, and the description of the latter species should, there-
fore, be consulted for data on the orbit of Parapontoporia.

The vomer forms a deep, narrow keel between the internal
nares, and is continuous with the very deep, narrow keel
formed by the palatines between the fossae for the pterygoid
sinuses (Fig. 20c). These narrow fossae are closely appressed
on either side of this very deep and narrow keel at the pos-

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Barnes: Pontoporiid Dolphins 15

Figure 9. Parapontoporia sternbergi (Gregory and Kellogg, 1927), referred periotics from the San Diego Formation: UCMP 88581, right: a,
cerebral view; b, tympanic view; UCMP 88589, left: c, cerebral view; d, tympanic view; LACM 58901, left: e, cerebral view; f, tympanic view;
LACM 103975, left: g, cerebral view; h, tympanic view. All natural size.

terior end of the palate, and they extend anteriorly only slight-
ly beyond the level of the antorbital notches. However, they
diverge dorsoposteriorly and become enlarged as they ascend
into the skull anterior to the nares (Fig. 8b). Such a dorsal
expansion of the pterygoid sinus is not common in odon-
tocetes, but does occur also in Lipotes vexillifer and Ponto-
poria blainvillei.

The palate is fairly flat for most of its length; not quite as
flat as in Pontoporia blainvillei, and more deeply fissured
medially at the posterior end. Premaxillae appear about mid-
length on the palate between the maxillae, from which they
are separated by elongate sutures. Progressing anteriorly, the
premaxillae occupy increasing amounts of the palatal surface
so that near the rostral extremity the medial side of the al-
veolar row is composed entirely of premaxilla (Fig. 8b). The
vomer is exposed on the posterior one-fourth of the palate.

On the posterior part of the rostrum the alveolar rows
curve slightly dorsally. Here the surface of the maxilla be-
comes increasingly porous, except for a band of relatively
dense bone extending along the dorsal edge of the alveolar
row. The alveoli for the teeth are all approximately 2 mm
in diameter. The deepest part of the rostrum is at the prox-
imal end ventral to the antorbital notches. Here it is trian-
gular in cross section with a prominent ventral keel. Anterior
to this keel are a pair of palatine foramina in each maxilla
on either side of the midline. At the deepest part of the keel
lie the triangular palatines, wedged between the maxillae.

Periotic. I have previously (Barnes, 1973a:figs. 2a, b) il-
lustrated a periotic (UCMP 57991) from the San Joaquin
Formation, a correlative of the San Diego Formation, and
identified it as “ Stenodelphis ” sternbergi. Periotics like this
one have been reported from the San Diego Formation

(Barnes, 1973a; Demere, 1981:fig. 5) and more are described
here, but none has ever been found in association with a skull
of P. sternbergi nor of any other species of Parapontoporia.

I now assign seven periotics collected from the San Diego
Formation to P. sternbergi based upon such resemblances to
periotics of Lipotes vexillifer as: comparable proportions
among the different parts of the periotic, pointed anterior
process with a separate rugose process on its medial side, a
curved crest that extends along the lateral side of the anterior
process from its tip to a tuberosity immediately anterior to
the groove for the tympanic membrane, a relatively small,
slightly concave and grooved articular surface for the bulla
on the posterior process, a deep cleft between the anterior
process and the cochlear portion of the periotic, and a rel-
atively small, circular internal acoustic meatus. The periotic
of Lipotes vexillifer (see Brownell and Herald, 1972:pl. 3,
figs. 1-6; Kasuya, 1973:pl. 10, figs. 1 6—20; Zhou, Qian, and
Li, 1979:pl. 3, figs. 1-6) differs from that of P. sternbergi by
being much larger and by having a relatively larger cochlear
portion with more rugosities on its cerebral surface and a
more circular internal acoustic meatus.

The periotic of Pontoporia blainvillei (see Kasuya, 1973:
pi. 9, figs. 17, 20-23) is considerably different from those of
both of the above species by being very small and by having
relatively smaller anterior and posterior processes, a smaller
tuberosity anterior to the groove for the tympanic membrane,
a more inflated cerebral surface, a distinctive reticulated,
etched pattern on the tympanic side of the cochlear portion,
and by lacking a process on the medial side of the anterior
process.

The periotics of P. sternbergi from the San Diego For-
mation are variable (Fig. 9) in the shape and degree of infla-

16 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

Figure 10. Parapontoporia sternbergi (Gregory and Kellogg, 1927), referred specimens from the San Diego Formation: UCMP 88587, mandible
fragment: a, dorsal view; b, left lateral view; c, cross section; UCMP 88588, premaxillary fragment with teeth: d, lingua! view; e, posterior
view of the complete tooth. Scale line is for Figs, d, e only. Figs, a-c natural size.

tion of the cochlear part and the anterior process, and in the
size of the posterior facet for the tympanic bulla.

Mandible. The holotype is a section of a mandible, not a
rostrum as it was identified by Gregory and Kellogg (1927).
The specimen has no evidence of a vomer or mesorostra!
gutter which would be present if it were a rostrum, and it is
fused at the midline, which is the mandibular symphysis. All
skulls of P. sternbergi, in a manner typical of odontocetes,
exhibit no fusion on the palatal surfaces between the opposite
premaxillae or maxillae.

The mandible of P. sternbergi is long and slender, and the
dentaries are fused for most of their tooth-bearing length by
a firm suture. One deep, longitudinal groove extends along
the side of each dentary just below mid-height (Fig. 10c).
The dorsal margin of this groove projects ventrally so that
the opening of the groove is slightly constricted. Anteriorly
directed nutrient foramina scattered along the dentary emerge
into the upper part of this groove. The groove is less distinct
at its posterior end at a point anterior to the end of the
symphysis. The ventral surface of the mandible has a broad
and flat (or in some specimens a slightly convex) surface and
is marked along its midline by a faint groove tracing the
position of the mandibular symphysis. The whole surface of

the mandible, including the inside of the longitudinal grooves,
is composed of dense, striated bone. Along the midline of
the dorsal surface of the symphyseal region of the mandible
there is a broad, shallow groove with a slight longitudinal
ridge that marks the midline and the position of the man-
dibular symphysis (Figs. 10a, c). There is no median groove
in the mandible of Pontoporia. Teeth are set 1 to 2 mm apart
in circular alveoli separated by septa of cancellous bone. The
alveoli are 2 to 3 mm in diameter.

Teeth. The teeth in the holotype mandible fragment
(AMNH 21905) are not typical of those in the sample of
specimens now available. Some sort of possibly postmortem
mechanical or chemical attrition has reduced the crowns of
all the teeth in AMNH 21905 to featureless, rounded cores
lacking any enamel (Gregory and Kellogg, 1927:fig. 1). The
roots of the teeth are more expanded anteroposteriorly, es-
pecially at their apices, than those in any other specimen yet
recovered from the San Diego Formation. This is the feature
which Gregory and Kellogg ( 1 927:3, fig. 3) described as being
like a battle-ax. Because the holotype is larger than the other
available mandible fragments, I attribute this exceptional
root development to extreme old age of the individual.

Upper and lower teeth are similar in morphology. An un-

Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins 17

worn tooth (Figs. lOd, e) has a crown that is slender, slightly
compressed anteroposteriorly, with a nearly straight labial
margin and a prominent lingual bulge located proximally
near the enamel margin. The enamel is smooth, having no
rugosities or crests. No specimen has a complete dentition,
but it can be seen in the partial dentitions in the sample
available that the more anterior teeth in both the skull and
mandible have relatively slender and high crowns, and that
progressing posteriorly they are thicker, shorter, and more
curved lingually. The anterior teeth are positioned vertically,
but the middle and posterior teeth tilt more labially.

The part of the root that is exposed between the bony
alveolus and the enamel line of the crown is encircled by a
collar which is most prominent anteriorly and posteriorly
forming “shoulders” on the root. This condition is preserved
on the teeth of the holotype as well. Roots are flattened
transversely, particularly so at their apices, which are curved
posteriorly (Fig. lOd).

The teeth, or their alveoli, in the nearly complete skull
LACM 6238 total at least 77 on each side, and the rostrum
was probably at least 20 mm longer when complete. Because
the number of teeth in P. sternbergi is so close to the known
number of 80 to 82 on each side of each jaw in the holotype
of P. pacifica, I believe that the two species had similar num-
bers of teeth in both the rostrum and mandible.

Parapontoporia wilsoni , new species

Figures 11-16, 20b, 21a

Stenodelphininae, genus and species new (part): Barnes, 1977:

331.

DIAGNOSIS OF SPECIES. A species of Parapontoporia
differing from P. pacifica by having skull with premaxillae
depressed, forming a basin on proximal surface of rostrum
anterior to level of antorbital notches; differing from P. stern-
bergi by having facial region longer than wide (antorbital
notch to occipital crest versus supraorbital width), deeper
basin on proximal surface of rostrum, antorbital process
smaller, cranial vertex (comprised of frontals and nasals)
lower and less compressed transversely, nares not as verti-
cally oriented in passage through the skull but curving more
around anterior wall of braincase, temporal fossa more elon-
gate anteroposteriorly, zygomatic process of squamosal less
inclined anteriorly, squamosal between postglenoid process
and paroccipital process not as greatly compressed antero-
posteriorly; and differing from both P. sternbergi and P. pa-
cifica by having shorter fossae for pterygoid sinuses on pal-
atines.

HOLOTYPE. UCMP 83790, incomplete skull consisting
of the facial portion of the braincase, the posterior part of
the rostrum and the basicranium, missing part of basioccip-
ital, supraoccipital, squamosals, pterygoids, parietals, and
basisphenoid, collected by John Stanley prior to 1966.

TYPE LOCALITY. UCMP V-6969, in the sea cliff north
of Manresa Beach, Santa Cruz County, California.

FORMATION AND AGE. The lower part of the Purisima
Formation, latest Miocene, correlated with the “Jacalitos”
provisional mega-invertebrate stage of Addicott (1972) and

indirectly with the Hemphillian North American land mam-
mal age, and approximately 6 to 8 million years old. The
type locality at Manresa Beach is a considerable, but as yet
undetermined distance higher stratigraphically than the base
of the Purisima Formation and the type locality of the pin-
niped Dusignathus santacruzensis Kellogg, 1927. The age of
the lower part of the Purisima Formation has been considered
by Cummings, Touring, and Brabb ( 1 962) as correlative with
the Jacalitos Formation in the San Joaquin Valley, and by
Barnes (1977) and Repenning and Tedford (1977) as also
correlative with the lower member of the Almejas Formation
on Isla Cedros, Baja California, Mexico.

ETYMOLOGY. The species name honors the late Mr.
Leslie E. Wilson, who was a teacher, a collector, and re-
searcher of fossil odontocetes, and a generous benefactor to
the University of California Museum of Paleontology. Mr.
Wilson helped establish the Remington Kellogg Memorial
Fund at the Museum to support student research on fossil
marine mammals.

DESCRIPTION. The holotype of P. wilsoni (Figs. 11-16)
may be compared with the entire cranium and the proximal
part of the rostrum of P. sternbergi , but only the anterior
part of its facial region and the proximal part of its rostrum
are directly comparable with the holotype of P. pacifica. To
avoid the repetition that would arise from describing iden-
tical structures known for the two previously described species
of Parapontoporia. those structures that are not known in the
others, or those that serve to differentiate P. wilsoni from
them, will be emphasized in the following text.

The braincase of P. wilsoni is more elongate anteropos-
teriorly than that of P. sternbergi (Fig. 21). In P. wilsoni the
facial region is relatively longer, the cranial vertex is not so
steeply peaked, the nares do not pass so steeply into the skull,
the temporal fossa is longer anteroposteriorly, and the pos-
terior part of the squamosal, above the mastoid region and
between the glenoid fossa and the paroccipital process, is not
so compressed in an anteroposterior plane.

The basin that is located in the center of the proximal part
of the rostrum is approximately 8 mm deep; nearly twice as
deep as it is in P. sternbergi. In P. pacifica, there is no such
basin (Figs. 20a-c). The premaxillary foramina are located
approximately 4 mm farther posterior, relative to the antor-
bital notches, than they are in P. sternbergi. Each foramen
has the three typical sulci connected with it (anteromedial,
posteromedial, and posterolateral), however, the anterior side
of the premaxillary foramen and the proximal part of the
anteromedial sulcus are roofed over by bone (Fig. 1 1 ), even
more so than in P. sternbergi, in an unusual manner. The
posterolateral sulcus is very deeply incised into the premax-
illa and its lateral edge is overhung by a sharp lip of bone.
Posterior to this, the posterolateral sulcus curves along the
lateral edge of the premaxilla and becomes indistinct opposite
the posterior edge of the nares.

The posterior ends of the premaxillae are complete on the
holotype of P. wilsoni, and confirm the previously given de-
scription of this area in P. sternbergi. The right spiracular
plate is wider than the left, and the posterior end of the left
premaxilla extends farther posteriorly than the right. Both

18 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

premaxillae extend posteriorly closer to the anterolateral cor-
ners of the nasals than they do in P. sternbergi, and this I
interpret as a more primitive character.

The supraorbital region is more complete on the holotype
of P. wilsoni than it is on any specimens of the other species
of Parapontoporia, and the antorbital processes and maxil-
lary crests are perfectly preserved. Each antorbital notch is
approximately 8 mm deep and, in addition to being partly
constricted medially by the maxillary eminence, is partially
overlapped laterally by the anterior end of the anteromedially
canted maxillary crest on the supraorbital process. The max-
illary crests are similar in location, but relatively smaller than
those in Lipotes vexillifer, Pliopontos littoralis, and Ponto-
poria blainvillei. In Parapontoporia wilsoni, the apex of the
crest forms an uninterrupted arc from the antorbital process
to the postorbital process. Its lateral surface is slightly convex
and comprised of maxilla as well as of the frontal and lacrimal
above the orbit. In Pontoporia this crest is similarly shaped,
but is higher, narrower, located closer to the premaxilla, and
the lateral surface of the crest is rugose, excavated and slopes
more medially, and concomitantly, the frontal and lacrimal
are more exposed dorsally. The same crest in Lipotes vex-
illifer is developed into more of a knob.

In Parapontoporia wilsoni there is a wide fossa in the max-
illa medial to the maxillary crest that extends from the an-
torbital notch posterolaterally toward the postorbital process.
The large posterior maxillary foramen is located medial to
this depression where the maxillary surface changes its slope
and ascends toward the narial region and the cranial vertex.

Compared with P. sternbergi, P. wilsoni has the following
primitive characters. The occipital condyles protrude more
prominently from the occipital shield (Figs. 1 3, 14), the pos-
terior part of the temporal fossa is not roofed over as far
laterally by the maxilla and frontal, the paroccipital process
is not located as far anteriorly on the braincase and therefore
is located less beneath the temporal fossa, and the posterior
part of the zygomatic process of the squamosal is not com-
pressed against the paroccipital process and therefore the
stemomastoid muscle fossa and the intervening bone that is
dorsal to the mastoid process is nearly three times the width.
The falcate processes of the basioccipital are thinner and are
more excavated laterally for the peribullary sinus than in P.
sternbergi. This is a derived character. On the holotype of P.
wilsoni, the falcate process is in many places only 1 to 2 mm
thick, and in one place it is so thin that, even in its fossilized
state, it will transmit light. The zygomatic process of the
squamosal is farther from the braincase wall and the inter-
vening squamosal fossa is therefore wider. The shelf of bone
anterior to this fossa is small and not upturned as in P.
sternbergi. The lateral margin of the glenoid fossa forms a
more prominent, vertical border (Figs. 15, 16).

The orbit is small and the frontal forms the posterior two-
thirds of its roof. The lacrimal is large compared with species
of Delphinidae and forms the anterior one-third of the orbital
roof (Fig. 16). In Pontoporia blainvillei and Lipotes vexillifer
the lacrimal is larger yet.

The anterior end of the jugal, as is typical of most modem
odontocetes, is fused to the lacrimal, extends anteriorly be-

neath the maxillary eminence on the proximal part of the
rostrum and is surrounded dorsally and ventrally by the max-
illa. The zygomatic process of the jugal departs from the body
of the fused jugal and lacrimal on the anteromedial side of
the antorbital notch beneath this eminence. This location of
departure of the zygomatic process is similar to that in Li-
potes vexillifer, but different from that in Pontoporia blain-
villei in which it is located posterior to the antorbital notch.
No known specimen of Parapontoporia has a complete jugal.

The optic foramen leaves the braincase beneath a relatively
massive strut of bone on the frontal lying posterolateral to
the naris. The orbital apertures of the infraorbital foramen
complex join to form a large recess in the medial wall of the
orbit. These foramina connect dorsally with the maxillary
and premaxillary foramina. There is a large but shallow fossa
under the anterior part of the supraorbital process indicating
that this area held a preorbital lobe of the pterygoid sinus of
the middle ear air sinus system.

PHYLOGENETIC RELATIONSHIPS

Before discussing interrelationships of the subfamilies within
the family Pontoporiidae, it is important to clarify which
previously reputed pontoporiids are not, in my opinion, de-
monstrably members of the group. Lonchodelphis occiduus
(Leidy, 1 868) is a problematical dolphin from the latest Mio-
cene Purisima Formation in central California. The species
was based upon a rostral fragment, and was believed by Allen
(1924) to be related to Pontoporia after it had been allied
first with Delphinus by Leidy ( 1 868) and later with Phocoena
by Jordan and Gilbert (1919). I have (Barnes, 1977:332;
1 984: 1 2) pointed out that the specimen is not similar to any
specimen of a pontoporiid. No other specimens have ever
been assigned to the species and, until a more complete spec-
imen is identified, it should be considered as some unknown
type of odontocete. Simpson ( 1 945: 1 0 1 ) classified in the Sten-
odelphminae the Pliocene age Argentinian species, Pontivaga
fscheri Ameghino, 1891, which was based on a lower jaw
fragment. The species cannot be objectively compared with
another, contemporaneous pontoporiid from Argentina,
Pontistes rectifrons Burmeister, 1885, however, which was
based on a skull (see Barnes, 1984: 11, 12). de Muizon (1983)
suggested that the Late Pliocene age Peruvian species, Plio-
pontos littoralis was very closely related to Recent Pontoporia
blainvillei and might have evolved from Pontistes rectifrons.
I accept the idea that Pontoporia, Pontistes, and Pliopontos
are related, and Pontivaga must be considered as an odon-
tocete of uncertain affinities until a mandible of Pontistes
rectifrons has been compared with it.

In the classification I propose here, the family Pontopo-
riidae includes three subfamilies. The nominate subfamily
Pontoporiinae includes Recent Pontoporia blainvillei, the
closely related Pliocene Peruvian Pliopontos littoralis de
Muizon, 1983, and the Pliocene Argentinian species, Pon-
tistes rectifrons Burmeister, 1885. These species all have sym-
metrical cranial vertices. Parapontoporia, the sole genus in
the subfamily Parapontoporiinae, has teeth, rostrum, and
some cranial characters like Pontoporia, but has an asym-

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Barnes: Pontoporiid Dolphins 19

20 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

Figure 11. Parapontoporia wilsoni, new species, holotype, UCMP 83790, partial skull, dorsal view, natural size.

aon

Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins 21

Figure 12. Parapontoporia wilsoni, new species, restoration of partial skull based on holotype, UCMP 83790. dorsal view, natural size.

Figure 13. Parapontoporia wilsoni, new species, holotype, UCMP 83790, partial skull, left lateral view, natural size.

OUI

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Barnes: Pontoporiid Dolphins 23

Figure 14. Parapontoporia wilsoni, new species, restoration of partial skull based on holotype, UCMP 83790, left lateral view, natural size.

24 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

Figure 15. Parapontoporia wilsoni, new species, holotype, UCMP 83790, partial skull, ventral view, natural size.

dod

Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins 25

Figure 16. Parapontoporia wilsoni, new species, restoration of partial skull based on holotype, UCMP 83790, ventral view, natural size.

a

b

Figure 17. Skulls of Pontoporiidae: a, Pontoporia blainvillei (Gervais and d’Orbigny, 1844); b, Parapontoporia sternbergi (Gregory and
Kellogg, 1927); c, Lipotes vexillifer Miller, 1918. (a after Flower, 1869:pl. 28, fig. 3; b after Fig. 7b, this paper; c after Brownell and Herald,
1972:fig. 1.)

metrical cranial vertex and some other derived cranial char-
acters like Lipotes. Parapontoporia, therefore, is in many
ways intermediate between Pontoporia and Lipotes. The lat-
ter was designated by Zhou, Qian, and Li (1979) as the type
genus of a new family, the Lipotidae. To reflect the close
relationships and to balance the classification, I recognize the
Lipotinae as a third subfamily of Pontoporiidae.

Zhou, Zhou, and Zhao (1984) described a fossil that is

possibly Miocene in age as an extinct member of the Lipo-
tidae, calling it Prolipotes yujiangensis. All that is known of
the species is the holotype mandible fragment, but the spec-
imen does have morphology very similar to the extant Li-
potes vexillifer. Odontocete mandibles generally have fewer
diagnostic characters than do skulls, and as a rule it is unwise
to make taxonomic inferences from them. There is nothing
in the morphology of P. yujiangensis, however, that would

26 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

preclude a possible relationship between it and L. vexillifer,
and I provisionally classify both species in the same subfam-
ily.

In summary, the family Pontoporiidae includes three
subfamilies: the Pontoporiinae containing Pontoporia blain-
villei and extinct taxa ( Pontistes rectifrons, Pliopontos litto-
ralis) that have symmetrical crania, very long rostra, and
extreme polydonty; the Parapontoporiinae including species
of Parapontoporia, which have asymmetrical crania, very
long rostra, and extreme polydonty; and the Lipotinae in-
cluding Lipotes vexillifer, which has an asymmetrical cra-
nium, shorter rostrum, and only moderate polydonty (Fig.
17). The fossil Prolipotes yujiangensis may belong to the
latter subfamily, but no cranial material is known that could
confirm this. The family Pontoporiidae is united by a unique
suite of shared derived characters as given in the family
diagnosis.

The deep grooves on the lateral sides of the dentaries in
fossil Parapontoporia sternbergi and Recent Pontoporia
blainvillei are convergent derived characters (autapomor-
phies). I base this conclusion on two lines of evidence. (1)
The most primitive, and chronologically oldest species of the
genus Parapontoporia, P. pacifica, has no mandibular grooves.
Instead it has a shallow longitudinal sulcus on each dentary.
Deep mandibular grooves are present, however, in the chron-
ologically youngest species, P. sternbergi (Figs. 20d, e). (2)
Each dentary of Recent Pontoporia blainvillei has only one
mental foramen in the mandibular groove, and this enters
the groove at its posterior end. In Parapontoporia each den-
tary has four or five foramina that are spaced out along the
length of the mandible. This is the case in species of Para-
pontoporia both with and without a mandibular groove, and
is the primitive (plesiomorphic) condition among odonto-
cetes.

In Pontoporia blainvillei, and apparently also in Parapon-
toporia sternbergi, the nerves and blood vessels emerging
from the mental foramina would lie in these grooves, and
presumably derive some measure of protection from them.
Therefore, the function of the mandibular grooves is the same
in both genera, but their origins are separate.

I believe that the unique features of the teeth of Pontoporia
blainvillei and Parapontoporia spp. are shared derived (syn-
apomorphic) characters because even the earliest species of
Parapontoporia, P. pacifica, has them. The rugose texture of
the enamel on the teeth of Lipotes vexillifer and Prolipotes
yujiangensis is probably a primitive character. Many prim-
itive odontocetes have rugose enamel, and this I regard as a
carryover from earlier squalodonts and agorophiids. Most
modem species have smooth enamel, the derived condition.
Inia geoffrensis, often classified with Lipotes vexillifer in ear-
lier works, has rugose enamel, but the posterior teeth have
very large lingual shelves and the dentition is otherwise not
very much like that of Lipotes vexillifer. Teeth of both Lipotes
vexillifer and Prolipotes yujiangensis have a slightly swollen
shoulder on the root, and this is a derived character that is
shared with Pontoporia blainvillei and Parapontoporia spp.

An asymmetrical cranial vertex that is offset to the left
side is a convergent derived character that has appeared sep-

arately in different groups of odontocetes at different times.
Prior to Late Miocene time, representatives of most odon-
tocete families, with the exception of such groups as sperm
whales (Physeteridae) and beaked whales (Ziphiidae), had
symmetrical cranial vertices and narial regions. Lipotes vex-
illifer and Parapontoporia spp. have, among other shared,
derived cranial characters, asymmetrical cranial vertices and
asymmetrical bones around the external nares. The cranial
vertices of Pontoporia blainvillei and the fossil species of
Pontoporiinae, however, are symmetrical, as well as being
lower and more elongate anteroposteriorly than those of
Parapontoporia spp. or Lipotes vexillifer. Because Pontoporia
blainvillei has asymmetrical nasal diverticula (Schenkken,
1972) and because so many of its other features are derived,
I suspect that its cranial symmetry is not primitive, but is a
reversal from an earlier asymmetrical condition.

If the above arguments are accepted as valid and Ponto-
poria, Parapontoporia , and Lipotes are indeed closely related,
then there are at least two possible diagrams that would show
their interrelationships, depending on which of the above
characters are considered to be shared and derived versus
convergent and derived.

In the most parsimonious scheme, and the one that I prefer
(Fig. 18), the similar structures of the teeth of Parapontoporia
and Pontoporia are interpreted as shared derived characters
(synapomorphies) and the deep mandibular grooves as con-
vergent derived characters (autapomoprhies). The asym-
metrical cranial vertex of Parapontoporia spp. and Lipotes
vexillifer, while being a derived character relative to more
primitive odontocetes, is the shared primitive character state
(symplesiomorphy) for Pontoporiidae, and the symmetrical
cranial vertex of Pontoporia (and fossil Pontoporiinae) is a
unique derived character (autapomorphy), being secondarily
symmetrical and a reversal from the asymmetrical condition.

Another possible interpretation of relationships (Fig. 19)
is one in which the symmetrical cranial vertex of Pontoporia
blainvillei is considered to be primitive for the family, and
the asymmetrical vertices of Parapontoporia spp. and Lipotes
vexillifer are shared derived characters. This is more consis-
tent with traditional ideas about acquisition of cranial asym-
metry in odontocete families, but requires an assumption
that in addition to deep mandibular grooves being conver-
gent, as in the first case above, that the unusual Pontoporia-
like teeth and the very long rostrum are the primitive char-
acter state for Pontoporiidae. In that case the lesser tooth
count, thick, short rostrum and mandible, and rugose tooth
enamel of Lipotes vexillifer are derived characters and all of
these would then necessarily be interpreted as secondary re-
versals back to the primitive odontocete condition. In each
of the two possible schemes of relationships discussed above,
the intermediate position of Parapontoporia spp. and the
polarity of characters and inferred interrelationships of its
included species remain unchanged.

Within the genus Parapontoporia, the most primitive
species is Late Miocene P. pacifica, with no mandibular
grooves and no rostral basin (Figs. 20a, d). The approxi-
mately contemporaneous species, P. wilsoni, however, is more
derived and has a very deep rostral basin (apomorphy), but

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Barnes: Pontoporiid Dolphins 27

Figure 18. Diagram of the most parsimonious scheme of relationships among taxa of Pontoporiidae, with the fewest implied convergences
and evolutionary reversals, but requiring the assumption that the cranial symmetry of Pontoporia blainvillei is the result of a secondary reversal
from an earlier asymmetrical condition, a contradiction to traditional ideas regarding odontocete evolution. Characters marking the dichotomies
are as follows: (1) The primitive state for Odontoceti is a symmetrical cranial vertex. (2) An asymmetrical cranial vertex (apomorphy) is
presumed to be the primitive character state of the earliest members of the Pontoporiidae. The family is differentiated from all other groups
of Odontoceti, including all other families within the superfamily Platanistoidea, by the shared characters given in the family diagnosis.
Interrelationships of the other platanistoid families are insufficiently understood, within the context of the analysis presented here, to be shown
separately in this scheme. (3) Lipotes vexillifer retains a relatively short, thick rostrum and mandible, and rugose enamel on the teeth as
primitive characters (subfamily Lipotinae, presumably including Prolipotes yujiangensis, but the species is insufficiently known to include on
the diagram). (4) Parapontoporia and Pontoporia share as synapomorphies: polydonty, Pontoporia- like teeth, extremely long rostrum and
mandible, and a bony wall in the orbit formed by a posterior extension of the lateral lamina of the pterygoid and, at least in part, also by
posterior extensions of the palatine and maxilla. These characters are shared by the subfamilies Parapontoporiinae and Pontoporiinae. (5)
Exceptionally long rostrum, extreme polydonty (80-82 teeth in each side of each jaw in contrast with 48-61 in Pontoporia blainvillei) are
autapomorphies for the genus Parapontoporia (subfamily Parapontoporiinae). (6) The most primitive species in the genus. (7) Basin formed
on the proximal part of the rostrum; a derived character shared by P. wilsoni and P. sternbergi. (8) Very deep rostral basin (autapomoprhy),
otherwise more primitive than P. sternbergi, especially in having a braincase that is less foreshortened anteroposteriorly (less telescoped). (9)
Anteropostenorly compressed (more telescoped) cranium, more vertical narial passages, more elevated and more transversely compressed
cranial vertex, and deep mandibular grooves are autapomorphies of P. sternbergi. The rostral basin is shallower than in P. wilsoni, and in
this regard P. sternbergi is more primitive. (10) Pontoporia blainvillei has the following autapomorphies: vomer not exposed on palate,
spiracular plates convex and elevated, posterior premaxillary terminations shortened, shallow squamosal fossa between zygomatic process of
squamosal and braincase. Where preserved, these characters are also present in the fossil species, Pontistes rectifrons and Pliopontos littoralis,
but the scope of this study does not include a more detailed analysis of the relationships of the latter species (subfamily Pontoporiinae).

a mandible has not been found and it is not known whether
or not it had lateral grooves (Fig. 20b). These two species
represent two lines of descent within the genus. The much
younger species, the Late Pliocene P. sternbergi, has deep
mandibular grooves (apomorphy), but only a shallow rostral
basin (Figs. 20c, e). Morphologically and temporally it could

have evolved from P. pacifica. It would have had to have
undergone an evolutionary reversal, a shallowing of the ros-
tral basin, to have evolved from P. wilsoni. The braincase
of P. sternbergi is anteroposteriorly compressed (apomorphy)
when compared with that of P. wilsoni (Fig. 21). Because P.
pacifica is so primitive, it undoubtedly had an anteroposte-

28 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

Other

Odontoceti

Pontoporia

blainvillei

Parapontoporia

pacifica

Parapontoporia

wilsoni

Parapontoporia

sternbergi

Lipotes

vexillifer

Figure 19. An alternative, less parsimonious interpretation of relationships among taxa of Pontoporiidae. In this scheme the symmetrical
cranial vertex of Pontoporia blainvillei would be considered as the primitive character state for the family Pontoporiidae, as it is for primitive
odontocetes, and the asymmetrical cranial vertices of Parapontoporia spp. and Lipotes vexillifer would be synapomorphic. This arrangement
would suggest, however, that many other characters were the result of convergent evolution of derived states (e.g., long rostra, mandibular
grooves, polydonty, and the Pontoporia- like teeth in Pontoporia blainvillei and Parapontoporia spp.) or of evolutionary reversals back to
primitive states (e.g., short rostrum, low tooth count, and rugose enamel on teeth of Lipotes vexillifer).

riorly elongate braincase with proportions like that of P.
wilsoni. and if P. pacifica were ancestral to P. sternbergi, such
anteroposterior compression could plausibly have occurred
within the lineage in the elapsed time interval of several
millions of years.

CLASSIFICATION

Class Mammalia Linnaeus, 1758
Order Cetacea Brisson, 1762
Suborder Odontoceti Flower, 1867
Superfamily Platanistoidea (Gray, 1863) Simpson,
1945

Family Pontoporiidae (Gill, 1871) Kasuya, 1973
Subfamily Lipotinae (Zhou, Qian, and Li, 1979),
NEW RANK AND CONTEXT
Prolipotes Zhou, Zhou, and Zhao, 1984

Prolipotes yujiangensis Zhou, Zhou, and
Zhao, 1984. ?Miocene, China
Lipotes Miller, 1918

Lipotes vexillifer Miller, 1918. Recent, China

Subfamily Parapontoporiinae Barnes, 1984
Parapontoporia Barnes, 1984

Parapontoporia pacifica Barnes, 1984. Latest
Miocene, Baja California
Parapontoporia wilsoni, NEW SPECIES.

Latest Miocene, California
Parapontoporia sternbergi (Gregory and Kel-
logg, 1927). Late Pliocene, California
Subfamily Pontoporiinae (Gill, 1871) Barnes,
1984

Pontistes Burmeister, 1885
Pontistes rectifrons (Bravard, 1884). Plio-
cene, Argentina
Pliopontos de Muizon, 1983
Pliopontos littoralis de Muizon, 1983. Early
Pliocene, Peru
Pontoporia Gray , 1846

Pontoporia blainvillei (Gervais and d’Orbi-
gny, 1 844). Recent, Atlantic coast of South
America

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Barnes: Pontoporiid Dolphins 29

Pmx

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Barnes: Pontoporiid Dolphins

Figure 20. Comparisons of cross sections through the proximal part of the rostrum of a, Parapontoporia pacifica Barnes, 1 984; b, P. wilsoni, new species; c, P . sternbergi (Gregory
and Kellogg, 1927); and through mandibles at mid-length of the symphyseal area of d, P. pacifica; and e, P. sternbergi; all natural size.

Figure 21. Dorsal views of braincases of two species of Parapontoporia; a, P. wilsoni, new species, based on the holotype; b, P. sternbergi
(Gregory and Kellogg, 1927), based on referred specimens, principally LACM 6238; reduced to the same cranium length.

Use of either of two family group names, Pontoporiidae
(or Pontoporiinae) and Stenodelphinae (incorrectly emended
to Stenodelphininae), has varied among authors, depending,
in some cases, upon their acceptance or rejection of the no-
tion that Pontoporia Gray, 1846, is a valid senior synonym
of the genus name Stenodelphis d’Orbigny and Gervais, 1 847.

Especially during the early 1 900’s it was understood by many
authors that Pontoporia was a preoccupied name, and Steno-
delphis was the preferred usage. More recently, Pontoporia
has been confirmed as valid (Hershkovitz, 1961), and in
recent literature is the commonly used generic name.

The family group name based upon Pontoporia, Ponto-

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Barnes: Pontoporiid Dolphins 31

poriidae, is available from Gill's (1871) first use of Ponto-
poriinae as a monotypic subfamily of the Platanistidae. The
following year Gill (1872) reclassified the Pontoporiinae as
a subfamily in the family Delphinidae. Gray (1871) used the
incorrectly formed family name Pontoporiadae, which he
classified in the suborder Delphinoidea with the families In-
iidae, Delphinidae (including phocoenids), Grampidae,
Globicephalidae, Orcadae (sic) (these latter three are now
classified as synonyms of Delphinidae), and Belugidae (cor-
rectly called Monodontidae). True (1908) recognized the ge-
nus name Stenodelphis instead of Pontoporia, and therefore
proposed the subfamily Stenodelphinae which he classified
in the family Delphinidae. He was followed in this by Miller
(1923), Kellogg (1928), and many subsequent authors, in-
cluding myself (Barnes, 1977), who, however, employed the
incorrectly emended form of the name, Stenodelphininae.
Simpson (1945) used the same incorrectly formed name, but
classified the subfamily with the subfamilies Iniinae and Plat-
anistinae in the family Platanistidae. Rice (1967) recognized
the same hierarchy and ranks as did Simpson, but substituted
the earlier and correct name, Pontoporiinae. Kasuya (1973)
used Gill’s name at the family rank, Pontoporiidae, and he
classified the family in the superfamily Plantanistoidea, in
which he also included the families Iniidae (including Li-
potes) and Platanistidae. Zhou, Qian, and Li (1979) erected
the new monotypic family Lipotidae for Lipotes, which had
previously been classified in either Iniidae or Iniinae, and
Zhou (1982) classified Lipotidae, Iniidae, Pontoporiidae, and
Platanistidae as four separate families in the superfamily
Platanistoidea.

In summary, each of the living genera, Pontoporia, Lipotes,
Inia, and Platanista, has been the basis for establishment of
a family group name. Their ranks in published classifications
have varied between subfamily and family, and they have
usually been classified in the family Platanistidae or the su-
perfamily Platanistoidea correspondingly. Pontoporia has also
commonly been classified in Delphinidae, and Lipotes and
Inia have usually been classified together in the family Ini-
idae or subfamily Iniinae. In my classification Lipotes is
related to Pontoporia and classified in the Pontoporiidae, not
the Iniidae. Iniidae and Platanistidae are, therefore, separate
families.

CONCLUSIONS

The Parapontoporiinae are an extinct Late Miocene and Plio-
cene eastern North Pacific subfamily of the dolphin family
Pontoporiidae and are represented by one genus, Parapon-
toporia Barnes, 1984. This extremely long-snouted genus is
morphologically and zoogeographically intermediate be-
tween the living marine franciscana or La Plata dolphin,
Pontoporia blainvillei (Gervais and d’Orbigny, 1844), of the
southwest Atlantic and the living freshwater beiji or white
flag dolphin, Lipotes vexillifer Miller, 1918, of China. The
latter two are each placed in separate subfamilies of the Pon-
toporiidae, called the Pontoporiinae and Lipotinae, respec-
tively. There is a possible fossil relative of Lipotes Miller,
1918, Pro/ipotes vujiangensis Zhou, Zhou, and Zhao, 1984,
of questionable Miocene age from China. Two South Amer-

ican Pliocene fossil species, Pontistes rectifrons (Bravard,
1884) and Pliopontos littoralis de Muizon, 1983, are related
to Pontoporia. Lipotes had previously been classified in the
Iniidae, or more recently put in its own monotypic family,
Lipotidae.

The genus Parapontoporia has been documented previ-
ously by several published references to fossils from the lat-
itudes between approximately 27° and 38° north in California
and Baja California under such identifications as Stenodel-
phis (or “ Stenodelphis ”) sternbergi Gregory and Kellogg, 1 927.
Among these scattered records are at least three species, each
of which is characterized and diagnosed in the present study.

The oldest and most primitive of these is Parapontoporia
pacifica Barnes, 1984, from the latest Miocene age lower
member of the Almejas Formation on Isla Cedros, Baja Cal-
ifornia, and which is between approximately 6 and 8 million
years old. An approximately contemporaneous species, Par-
apontoporia wilsoni, new species, from low in the Purisima
Formation in central California differs from P. pacifica no-
tably by having a deep basin on the proximal surface of the
rostrum. A much younger species from the Late Pliocene age
lower member of the San Diego Formation at San Diego,
California, P. sternbergi (Gregory and Kellogg, 1927), is be-
tween approximately 2 and 4 million years old. This species
does not have as deep a rostral basin as P. wilsoni and has
a more anteroposteriorly compressed braincase. It also has
deep mandibular grooves that are not present in P. pacifica.
The species is the most abundantly represented of the three,
and is the only one known by a complete skull.

At least two schemes of interrelationships are possible be-
tween Lipotinae, Parapontoporiinae, and Pontoporiinae, each
of which would require the assumption that reversals have
occurred in the evolution of certain characters. The most
parsimonious hypothesis is that Lipotes is the most primi-
tive, that Pontoporia is the most derived, and that Parapon-
toporia is intermediate between them. This would indicate
that Pontoporia had secondarily acquired a symmetrical cra-
nial vertex, that the asymmetrical cranial vertices of Lipotes
and Parapontoporia are a shared primitive character for the
family and that the unusual teeth of Parapontoporia and
Pontoporia are shared and derived. In this case the rugose
enamel on the teeth and the shorter and thicker rostrum and
mandible of Lipotes would be primitive characters, and the
deep mandibular grooves of living Pontoporia blainvillei and
the Late Pliocene fossil Parapontoporia sternbergi are con-
vergent and derived.

ACKNOWLEDGMENTS

This study is derived directly from part of my Ph.D. thesis
which was submitted to the University of California at Berke-
ley, and I thank William A. Clemens, Donald E. Savage, J.
Wyatt Durham, and Frank C. Whitmore, Jr. for their advice
and comments during my studies. Preparation and illustra-
tion of most of the specimens was funded by the UCMP
through the Annie M. Alexander Endowment. Dick Meier
of LACM prepared the photographs, and Joe Nakanishi of
LACM helped in reproducing the artwork.

I benefited from many discussions with Robert L. Brow-

32 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

nell, Jr. regarding these dolphins, and I thank him, John M.
Harris, Samuel A. McLeod, Edward Mitchell, and David P.
Whistler for comments on the manuscript. Richard H. Ted-
ford and Michael O. Woodbume invited me to study the
fossil from Isla Cedros, and J. Howard Hutchison brought
to my attention the one from the Purisima Formation. I thank
Robert L. Brownell, Jr., Thomas A. Demere, Richard Eth-
eridge, Jason A. Lillegraven, James G. Mead, Donald Patten,
John Stanley, David P. Whistler, and the late Leslie E. Wilson
for either collecting or providing access to specimens used
in this study.

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Accepted 8 January 1985.

34 Contributions in Science, Number 363

Barnes: Pontoporiid Dolphins

IrafWknialsS WJwlir:' ; i'il'ii tin .l I , I i" . i.Vj,i:i ;.. i , - ■ ;

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'SI

!X

v

Number 364
13 June 1985

CONTRIBUTI1

MW*

! ■

j:

THE GEOLOGY OF THE RIO BENI: FURTHER EVIDENCE
FOR HOLOCENE FLOODING IN AMAZONIA

Kenneth E. Campbell, Jr., Carl David Frailey,

j risnsa. xjr a h.

and Jorge Arellano L.

iSS

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THE GEOLOGY OF THE RIO BENI: FURTHER EVIDENCE
FOR HOLOCENE FLOODING IN AMAZONIA

Kenneth E. Campbell, Jr.,1 Carl David Frailey,2 and
Jorge Arellano L.3

ABSTRACT. Quaternary sediments exposed in outcrops along the
Rio Beni, Bolivia, were deposited in two widely separated deposi-
tional realms. The Sierra Realm extends northeastward from the
front range of the Andes for 1 5-30 km, whereas the Monte Realm
encompasses all of the northernmost lowlands of Bolivia. In both
depositional realms the same stratigraphic sequence occurs: a lower
unit of clays with numerous channel-fill deposits at its base, capped
by two upper units of clayey silts and fine sands. The sediments of
the Sierra Realm were derived from the Bolivian Andes; by all
indications they were deposited contemporaneously with those of
the Monte Realm. The strata within the Monte Realm are correlated
with those to the northwest along the Rio Acre and Rio luma,
demonstrating that the deposits cropping out along the lower Rio
Beni represent the easternmost extension of the Inapari Formation,
a series of Holocene alluvial deposits derived from the Peruvian
Andes. The geological deposits of northern Bolivia provide sup-
porting evidence for the hypothesis of large-scale Holocene flooding
and deposition in southwestern Amazonia. A catastrophic flood re-
sulting from the sudden draining of glacial Lake Titicaca is proposed
to account for some of this flooding and several geological features
found in southwestern Amazonia.

RESUMEN. Los sedimentos cuatemarios expuestos en los estratos
a lo largo del rio Beni, Bolivia, fueron depositados por dos amplias
y separadas corrientes deposicionales. La “Corriente de la Sierra”
que se extiende al noreste, desde el frente de la Cordillera de los
Andes por 1 5 a 30 kilometros y la “Corriente del Monte” que abarca
la parte septentrional de las tierras bajas de Bolivia. En ambas cor-
rientes deposicionales se presenta la misma secuencia estratigrafica:
una unidad inferior de arcillas con numerosos canales basales y dos
unidades superiores compuestos por arcillas limosas y arenas finas.
Los sedimentos de la Corriente de la Sierra derivan de los Andes
bolivianos, por todas las evidencias ellos fueron depositados con-
temporaneamente a aquellos de la Corriente del Monte. Los estratos
de la Corriente del Monte son correlacionados con los descritos para
el rio Acre y rio Jurua al noroeste de Bolivia, de manera que los
depositos que se presentan aguas abajo del rio Beni representan la
extension mas oriental de la Formacion Inapari, una serie de de-
positos aluviales holocenicos derivados de los Andes peruanos. Los
depositos geologicos septentrionales de Bolivia son las evidencias
que sustentan la hipotesis de una inundacion a gran escala durante
el holoceno y la deposicion de sedimentos en la Amazonia meri-

dional. Una inundacion catastrofica resultante del repentino drenaje
del lago glacial Titicaca es propuesto para explicar varios rasgos
geologicos encontrados en el sudoeste de la Amazonia.

INTRODUCTION

The Amazon Basin and its neighboring lowlands have yet to
receive the in-depth, rigorous geological research warranted
a region so important to the South American continent, and
indeed, to the world. Occupying an area of over 1,800,000
km2, these vast lowlands are the subject of only a small
number of modem publications that discuss their geological
features and sediments (e.g., see citations in Petri and Fulfaro,
1 983). Almost all recent geological research has been directed
toward the discovery and exploitation of petroleum re-
sources; without recognized economic importance Quater-
nary geology has been very neglected. Aside from economics,
there are clear reasons for the lack of geological research in
Amazonia. The region is quite inaccessible, and river travel
is the only means of movement over vast areas. Furthermore,
the tropical forest, the largest such forest in the world, pro-
vides an almost complete cover of thick vegetation that limits
the study of surficial deposits to river cutbanks and well cores.
Nevertheless, geological research can and must be carried
out in these still remote lands in order that a solid core of
geological data is developed upon which future planned de-
velopments within the region can be based. This is partic-
ularly critical because preliminary data indicate that both

1. Section of Vertebrate Paleontology, Natural History Museum
of Los Angeles County, 900 Exposition Blvd., Los Angeles, Cali-
fornia 90007.

2. Department of Geology, Midland College, Midland, Texas
79705 and Research Associate in Vertebrate Paleontology, Natural
History Museum of Los Angeles County, Los Angeles, California
90007.

3. Servicio Geologico de Bolivia, Casilla 2729, La Paz, Bolivia.
Current address: Director, Instituto Nacional de Archaeologia, Casil-
la 5905, La Paz, Bolivia.

Contributions in Science, Number 364, pp. 1-18
Natural History Museum of Los Angeles County, 1985

ISSN 0459-8113

Figure 1 . Map of northern Bolivia showing the major outcrops along the Rio Beni, with distances in river-km (Servicio de Hidrografia Naval,
1 969) from Rurrenabaque: 1. Buena Vista (1 1); 2. Altamarani (17); 3. San Pablo (227); 4. San Roque (281); 5. Santa Catalina (345); 6. Guayaros
(355); 7. Candelaria (372); 8. Cavinas (385); 9. Carmen Alto (395); 10. Centro Navidad (410); 11. Barraca Rosario (421); 12. Fortaleza (437);
13. Ciudad California (465); 14. Etea (473); 15. Florencia (481); 16. Remanso (496); 17. Santa Elena (503); 18. Santa Rosa (510); 19. Sabacon
(515); 20. Mamorebey (544); 21. San Manuel (548); 22. San Martin (557); 23. Pena Amarilla (569); 24. Maracaibo (581); 25. Concepcion
(595); 26. San Miguel (601); 27. Brisa (621); 28. Barrio Lindo (627); 29. Palermo (694); 30. Candelaria (702); 31. Libertad (720); 32. Puerto
Gonzelo Moreno (728); Riberalta (758). The dashed line ( — ; lower left) indicates the northeastern limit of the front range of the Andes, while

the dotted line ( ; upper and lower right) indicates western limit of Brazilian Shield. Base map: Mapa Geologico de Bolivia, 1978, Yacimientos

Petroliferos Fiscales Bolivianos and Servicio Geologico de Bolivia, La Paz.

long-held assumptions (e.g., Abelson, 1983) and newly pop-
ular hypotheses (e.g., see papers in Duellman, 1979; Prance,
1982) concerning the late Cenozoic history of Amazonia re-
quire reevaluation.

The alluvial deposits covering the vast region of the Am-

azon Basin and its neighboring lowlands have always been
considered to be Pliocene to Pleistocene in age (e.g., Jenks,
1956; Ruegg, 1956; Kummel, 1948; ONERN, 1977; Petri
and Fulfaro, 1983; RADAMBRASIL, 1976, 1977). Recently,
however, Campbell and Frailey (1984, in press) have dem-

2 Contributions in Science, Number 364

Campbell, Frailey, and Arellano L.: Rio Beni Geology

onstrated that the alluvial deposits cropping out along the
Rio Acre in southwestern Amazonia and, by correlation,
similar deposits cropping out along the upper Rio Jurua about
500 km northwest of the Rio Acre, are Holocene in age.
Alluvial deposits forming three distinct units rest upon deep-
ly weathered Tertiary deposits along both of these rivers. A
characteristic basal clay-pebble conglomerate containing re-
worked Tertiary vertebrate fossils and fossil wood is a wide-
spread, but discontinuous, feature of these Holocene depos-
its. The deeply weathered Tertiary deposits are indicative of
late Pleistocene soil profiles, the upper portions of which were
removed prior to the deposition of the Holocene deposits.

To account for this blanket of Holocene sediments occur-
ring over such a broad area, Campbell and Frailey ( 1 984, in
press) proposed that massive flooding resulted from the melt-
ing of the Andean glaciers at the end of the Pleistocene. They
suggested that this flooding scoured the uppermost soil zones
developed on Tertiary strata in the region, depositing the
clay-pebble conglomerate in its wake. Further, they proposed
that this massive early flooding was followed by three distinct
cycles of deposition and erosion, the last period of erosion
being that which is underway today. Dates of about 5000 yr
B.P. and about 2800 yr B.P. were proposed to mark the ends
of the first two periods of deposition.

To determine if similar deposits of comparable age and
lithology occurred farther to the south, a geological survey
crossing the northern lowlands of Bolivia from the front range
of the Andes to the Brazilian Shield was executed via the
Rio Beni in July 1983. We reasoned that if three distinct
cycles of Holocene deposition and erosion had occurred over
wide areas of the Amazonian lowlands, evidence for these
cycles should exist in the alluvial deposits extending eastward
from the front range of the Andes. We conducted the survey
by means of boat travel from Rurrenabaque, at the base of
the front range (Fig. 1), to Riberalta, at the confluence of the
Rio Beni and Rio Madre de Dios. The transect was completed
by road between Riberalta and Guayaramerin, the latter a
town on the Bolivian-Brazilian border.

The survey was hampered by very unusual heavy rains
that fell in eastern Bolivia during the dry season of 1983, a
local expression of worldwide climatic anomalies occurring
at that time (Philander, 1 983a, b; Gill and Rasmusson, 1 983).
These rains maintained the level of the Rio Beni over 2 m
higher than what is normal for July, possibly preventing us
from observing the contact between the basal Quaternary
deposits and the underlying Tertiary strata. In other areas of

Figure 2. Satellite photo mosaic showing the course of the Rio
Beni from the Andes to Riberalta. The four regions of the river
discussed in the text are indicated. Northeast of the front range of
the Andes the light gray indicates areas slightly higher and better
drained than the areas of dark gray to the south. The light gray bands
bordering the rivers cutting through areas of dark gray indicate sig-
nificant natural levees. The white line running through the chain of
lakes points directly toward the point of exit of the main channel of
the Rio Beni from the front range of the Andes.

Contributions in Science, Number 364

Campbell, Frailey, and Arellano L.: Rio Beni Geology 3

southwestern Amazonia this contact, if visible at all, is seen
only when the rivers are at their lowest level during the dry
season. In addition, a newly deposited, deep layer of mud
made it very difficult, and in places impossible, to reach the
outcrops. All figures of outcrop height are relative to the level
of the river at the time we left Rurrenabaque, but unfortu-
nately this level could not be fixed relative to a known datum.
The fluctuating level of the river, which rose and fell over 1
m during the course of our trip as the consequence of a single
storm, endows all following figures of outcrop height with a
degree of error.

GEOLOGY

The eastern lowlands of Bolivia extend through the country
from its northern to its southern borders. The Andes flank
the lowlands on the west, with the front range in places rising
abruptly thousands of meters above the flat, swampy plains.
To the east the lowlands are bordered by the low sierras of
the Brazilian Shield, and many scattered remnants of the
Brazilian Shield stand isolated, surrounded by Quaternary
deposits. All of the rivers draining the northern two-thirds
of the Bolivian lowlands converge upon the Rio Madre de
Dios and exit Bolivian territory at its northeastern comer,
crossing over the crystalline rocks of the Brazilian Shield.
Because the terrain is so flat (e.g., the elevation of Rurrena-
baque is 227 m; Cobija, 202 m; Trinidad, 155 m; and Ri-
beralta, 135 m (Instituto Nacional de Estadistica, 1982), the
discharge from the rivers draining the eastern cordillera of
the Andes cannot be constrained to existing river channels
and the lowlands are consequently regularly inundated by
seasonal floods. (Note: the Servicio Nacional de Meteorolo-
gia de Bolivia (in Montes de Oca, 1983) gives the elevations
as: Rurrenabaque, 227 m; Cobija, 280 m; Trinidad, 236 m;
and Riberalta, 1 72 m.) The southern one-third of the eastern
Bolivian lowlands is, for the most part, an extensive area of
swamplands with no external drainage.

The geological deposits cropping out along the Rio Beni
were formed in two different depositional realms about 180
air-km apart. We consider a depositional realm to be a geo-
graphically well-defined area within which deposition of sed-
iments occurred during a specified time, in this case, the
Quaternary. The first depositional realm we crossed extends
northeastward from the front range of the Andes and is re-
ferred to hereafter as the Sierra Realm because of its prox-
imity to the mountains. The second depositional realm en-
compasses all of the northernmost Bolivian lowlands and is
referred to hereafter as the Monte Realm because of its thick
cover of tropical forest. The course of the Rio Beni east of
Rurrenabaque includes about 1 5 air-km in the Sierra Realm
and about 210 air-km in the Monte Realm. The two depo-
sitional realms stand out on satellite photographs (Figs. 2, 3)
as light gray areas, in contrast to the dark gray of the inter-
vening area which is lower in elevation. The differing shades
of gray result from different vegetation types; the lighter grays
indicate forests, the darker grays indicate wet savannas or
swamplands. The strips of light gray that cross the dark gray

areas, and within which lie the river channels, reflect the
higher topography of the natural levees bordering the rivers.

In the 180-air-km interval between the two depositional
realms only two small outcrops of possible Quaternary de-
posits were seen. Everywhere else in this interval the banks
of the Rio Beni are formed exclusively of Recent alluvium
with a relatively constant thickness of 2 m. At San Pablo
and San Roque (Fig. 1, Iocs. 3, 4) unstratified, variegated,
red, green, and gray clays extended to about 2-3 m above
the waterline. These clays are capped by about 1 m of Recent
alluvium that forms a flat terrace.

THE SIERRA REALM

The deposits of the Sierra Realm were all derived from the
Bolivian Andes immediately to the west. Included are the
alluvial fan deposits formed of sediments coming directly off
the eastern slopes of the front range and the deposits brought
from the valleys of the eastern Andes through the front range
by the Rio Beni and Rio Madidi.

There are only two outcrops along the Rio Beni within the
Sierra Realm that provide good sections: at Buena Vista and
Altamarani, 1 1 and 1 7 river-km, respectively, from Rurrena-
baque (Fig. 1 , Iocs. 1 , 2). The most complete section extends
from Altamarani downstream for about 1 km to just above
the confluence with the Rio Tuihuapa. About 9-10 m thick
at its highest, the section consists of three distinct units of
unconsolidated alluvial sediments (Fig. 4). The basal unit,
Member A, consists of poorly stratified gray, green, and yel-
low variegated clays about 2 m thick. Channel-fill conglom-
erates of small pebbles occur in the lower part of this unit.
A sharp contact separates these clays from the overlying unit
(Member B) of blocky, reddish clayey-silts about 3 m thick,
the top of which is marked by two prominent and other
smaller dark-gray paleosols. The uppermost unit. Member
C, consists of about 4-5 m of fine silts capped by a 20-30-
cm-thick layer of black soil.

The same three units occur in the Buena Vista section, but
Member C is much thinner, presumably through loss by
erosion. In addition, a cobblestone conglomerate passes un-
der Member A at the upriver end of the Buena Vista section.
Multiple paleosols are also present at the top of Member B
at Buena Vista.

Both the Buena Vista and Altamarani sections occur on
the north side of the river, and the terrain extending back
from the river’s edge is essentially flat. Except for one other
outcrop upriver from Buena Vista, also on the north side of
the river, the banks of the Rio Beni within the Sierra Realm
consists of Recent alluvium, about 2 m thick. The third
outcrop mentioned did not provide a clear section and was
only about 8 m thick. No clastic sediments coarser than sand
occur downriver from the Buena Vista section.

For the size of the area drained by the Rio Beni and its
tributaries, a mountainous region of approximately 67,000
km2 with peaks up to 7010 m elevation, the quantity of
Quaternary river deposits lying east of the front range is
remarkably small. Indeed, as can be noted on satellite pho-

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Campbell, Frailey, and Arellano L.: Rio Beni Geology

Figure 3. Satellite photo mosaic of northern Bolivia showing the extent of the Sierra and Monte Depositional Realms, the courses of the
major rivers, and the linear series of rectangular lakes extending northeastward from Rurrenabaque. The line of dots delimits the southern
extent of the Monte Depositional Realm and the line of triangles indicates the eastern limit of the Sierra Depositional Realm. From: Foto-
Mosaico Landsat de Bolivia, Edicion 1-P.E.B., 1975, Servicio Geologico de Bolivia. Reproduced by permission of the Servicio Geologico de
Bolivia.

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Campbell, Frailey, and Arellano L.: Rio Beni Geology 5

Figure 4. The geological section at Altamarani (Fig. 1, loc. 2) is
composed of three units. The lower unit (Member A) of channeled,
variably stratified clays is capped by two upper units (Members B
and C) of variably stratified, blocky silts, and fine sands. The two
dark bands at the top of Member B are prominent paleosols.

tographs (Figs. 2, 3), the eastward extent of the deposits left
by the Rio Beni at its point of exit from the front range
appears to be less than that of the alluvial fan deposits im-
mediately north and south of the river that represent depo-
sition of sediments derived solely from the eastern slopes of
the front range. We will return to this absence of expected
alluvial deposits later.

THE MONTE REALM

Trending north-northeastward, the Rio Beni cuts through the
Monte Realm from Santa Catalina (Fig. 1, loc. 5) to its con-
fluence with the Rio Madre de Dios at Riberalta. The total
areal extent of the Monte Realm is unknown, although from
satellite photographs (Fig. 3) it may be inferred to cover
approximately 90,000 km2 in Bolivia. The terrain immedi-

ately to the south of the Monte Realm is 1 5-20 m lower in
elevation, flat, swampy, and frequently inundated during the
annual rainy season. These areas of lower elevation appear
dark on the satellite photos (Figs. 2, 3), in contrast to the
lighter color of the better drained Monte Realm.

Outcrops are relatively common along the Rio Beni from
Santa Catalina to Riberalta, and the geologic sections are all
quite similar (see Fig. 1 for major localities). The generalized
section (Figs. 5, 6) is composed of three units of unconsoli-
dated alluvial sediments, each similar in lithology to the
corresponding unit seen in the Sierra Realm. The basal unit,
Member A, consists of about 3-5 m of variably stratified,
relatively pure, red, gray, green, and yellow variegated clays.
In the lower part of these clays occur many channel-fill de-
posits composed of fine-grained, well-sorted sands, silts, and
clay-ball conglomerates. The size of the multicolored clay
balls in the conglomerate range from less than 10 mm to
over 250 mm, and they occur both well sorted or poorly
sorted, and with or without a silt or sand matrix (Fig. 7a).

Layers of hematite 3-6 cm thick are a distinctive feature
of Member A. These occur between the channel deposits in
the lower part of the unit and also at the contact with silts
and sands of the overlying member. The layers are convo-
luted in many places (Fig. 8a, b), and represent groundwater
deposition of hematite at the contact between differing lith-
ologies.

Members B and C, each about 4-8 m thick, are similar
lithologically, being composed of silts and fine sands. In many
outcrops the contact between them may be readily distin-
guished only by sharp color differences (Fig. 5b), but at other
localities a distinct disconformity is visible (Figs. 5a, 6). At
some outcrops, such as at Ciudad California, channel de-
posits of sands and clay-ball conglomerates can be seen at
the top of Member B. When these channel deposits occur,
local layers of hematite may also be present as in Member
A, although they are usually much thinner.

The first complete section encountered in the Monte Realm
(Santa Catalina, Fig. 5a) differed from those farther down-

Figure 5. Two geological sections along the Rio Beni in the Monte Depositional Realm, a(left). Santa Catalina (Fig. 1, loc. 5). b. Cavinas
(Fig. 1 , loc. 8). Both sections show the hematitic concretionary zone in the B zone of the soil profile developed on Member C. The higher clay
content of the sediments at Santa Catalina is reflected in the surface texture of the outcrop.

6 Contributions in Science, Number 364

Campbell, Frailey, and Arellano L.: Rio Beni Geology

Figure 6. A partial view of the section at San Martin (Fig. 1, loc.
22) clearly shows the two contacts separating the three members of
the Inapari Formation. At this locality the base of Member B shows
good stratification.

river (e.g., Cavinas, Fig. 5b) by having a noticeably higher
clay content to Members B and C, and by being much thinner.
Both of these differences would be expected to occur at the
limit of alluvial deposition in what we have come to regard
as an environment of deposition similar to a broad delta.

The soil profiles visible in the river cutbanks of the Monte
Realm north of its southern limits vary in a consistent, pre-
dictable manner with the height of the outcrop, a direct re-
flection of the length of time the soil has had to develop. The
tops of the highest outcrops, here always at an elevation of
about 20 m above the level of the river, are taken to represent
the original surface formed by the depositional phase prior
to the onset of the present cycle of erosion. These outcrops
are everywhere characterized by a deep soil profile with a
dark red color. Included in this soil profile is a hematitic
concretionary zone about 0.5-1 m thick, lying about 2-3 m
below the ground surface (Fig. 5b). These hematite deposits
differ from the layers of hematite occurring lower in the
section at the contacts between different lithologies. Instead
of thick, solid layers, the hematite in the soil profiles occurs
as agglomerations of small (less than 1 cm diameter) con-

Figure 7. Representative clay-ball conglomerate that occurs in pa-
leochannels within Member B of the Inapari Formation, a. Variably
colored clay balls of various sizes with silt and sand matrix; on Rio
Beni at Ciudad California (Fig. 1, loc. 13). b. Unaltered clay balls
in well-sorted sand matrix; on Rio Acre (Fig. 1 ), just upstream from
the confluence of the Rio de los Patos. Scale in each figure equals
1 0 cm.

cretions that have in many places grown together. The dark
red color of the soil profile was most pronounced at the
concretionary zone.

This concretionary zone also appears in the thinner sec-
tions at the southern limit of the Monte Realm (Fig. 5a), but
here it is thinner and does not lie as deep below the surface
of the ground. This may reflect the higher clay content of the
sediments in this region. Throughout the Monte Realm the
A horizon of black humic material of the present soil profile
only locally reaches a thickness of more than 15-20 cm.

As in the Sierra Realm, at the major outcrops the terrain
extending back from the river’s edge usually appears flat, a
feature accentuated by the consistent placement of small set-
tlements or villages on these high, isolated reaches of land
with river access. The flat tops of the outcrops with incom-
plete sections indicate rivercut terraces, and within the Monte
Realm there are many discontinuous terraces at approxi-
mately the same elevations above the river. Two pronounced

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Campbell, Frailey, and Arellano L.: Rio Beni Geology 7

Figure 8. Thick layers of hematite separate Member A of the Ina-
pari Formation from overlying Member B. In a, the hematite layer
is shown as it appears in a roadside ditch near the Rio Buyuyo (Fig.
1), while in b the hematite layer is shown in cross section on the
Rio Beni near Barraca Rosario (Fig. 1, loc. 1 1). Note the similarity
in surface ribbing and texture. The thickness of the partially mud-
covered layer in b is 5-6 cm. This hematite layer was observed at
almost every outcrop within the Monte Realm where the contact
between Members A and B was exposed. Scale equals 1 5 cm.

terrace levels occurring at several localities along the river
are at about 9 and 1 5 m above the river. In addition to their
different elevations, these two terrace levels differ in the soil
profiles developed on them. At the 15 m level there is a 1-
2 m thick, dark-red soil horizon similar to that seen at the
top of the 20 m level, but no hematitic concretionary zone
was present. The red color of the B horizon gradually faded
into the C horizon and the original colors of the fine silts and
sands of Member B or C. The soil profile developed at the
9 m level did not have as dark a red B horizon, nor was it
as deep.

The lowest widespread terrace level north of the Rio Madi-
di was at about 3-4 m. This low terrace consisted of an upper

Figure 9. A river terrace at about 3-4 m above the level of the
Rio Beni. Above, showing cluster of homes on flat terrain; below,
showing the 2-3 m of Member A of the Inapari Formation capped
by about 1 m of Recent alluvium. The dark zone at the base of the
section reflects the dropping level of the river.

1 m cap of dark silts and sands representing Recent flood
deposits and a lower 2-3 m of the variegated red and yellow
clays of Member A, into which this terrace had been cut (Fig.
9).

The terrain between Riberalta, on the Rio Beni, and Gua-
yaramerin, on the Rio Mamore at the border with Brazil, is
typical of that of a flat plain dissected by a developing den-
dritic drainage system. This terrain continues right up to the
Rio Mamore, but directly across the river loom the north-
ernmost outlying hills of the Sierra dos Parecis, the western-
most range of the Brazilian Shield. Clasts of Shield rocks
were found in deposits in Bolivia west of Guayaramerin, but
they were not noted in deposits as far west as Riberalta.

CORRELATIONS

The outcrops along the Rio Beni can be correlated with those
that occur along the Rio Acre (Campbell and Frailey, 1984,
in press) (Fig. 10). The standard stratigraphic sequence (Fig.
11a), consisting of a basal unit of variably stratified clays
with channel-fill deposits of sands and clay-ball conglom-
erates (Fig. 7b) capped by two thicker units of relatively
unstratified, blocky silts and fine sands, is the same along
each river. Similarly the three units are separated by discon-
formities, although along the Rio Beni the contact between
the upper two units is not always as clear as it is along the
Rio Acre. Paleochannels occur at the contact between Mem-
bers B and C in some geologic sections of each river. Un-

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Campbell, Frailey, and Arellano L.: Rio Beni Geology

Figure 10. Two geological sections along the Rio Acre, demonstrating their similarity to those seen along the Rio Beni. a. Outcrop at San
Lorenzo, about 30 air-km upstream from Inapari, Peru, showing the various levels of the Inapari Formation. Tertiary clays are exposed in
the foreground. Member A of the Inapari Formation is covered, but the contact between Members B and C is clear, as is the hematite
concretionary zone near the top of Member C. This concretionary zone correlates with that seen in Fig. 5a, b. b. Partial section about 50 air-
km upstream from Inapari, Peru. The three members of the Inapari Formation are clearly seen, as is the contact with the underlying Tertiary
strata. The visible layer of hematite occurring at the contact between Members A and B of the Inapari Formation correlates with that seen in
Fig. 8a, b.

fortunately, along the Rio Beni the contact between the basal
clay unit and the underlying strata could not be seen, so it
is not possible to say if the characteristic basal clay-pebble
conglomerate occurring along the Rio Acre, Rio Buyuyo, Rio
Tahuamanu (Fig. 1), and Rio Jurua also occurs along the Rio
Beni.

The three lithologic units observed along the Rio Acre were
referred by Campbell and Frailey (1984, in press) to the
Inapari Formation of ONERN (1977). Radiocarbon dates
from four samples of wood from Member A of the Inapari
Formation taken at three different localities on the Rio Acre
ranged from 10,075 ± 150 yr B.P. to 5575 ± 105 yr B.P.
(Campbell and Frailey, 1984, in press). We here include the
Shiringayoc Formation of ONERN (1977) in the Inapari
Formation. The former was said to occur only along the
banks of the rivers, with the latter occurring in broad areas
between the rivers in the eastern part of the Departamento
de Madre de Dios, Peru, on the frontier with Bolivia. Our
field data indicate that the two formations are just one.

The hematite deposits observed along the Rio Beni were

not noted as a particularly significant feature along the Rio
Acre at the time of the fieldwork there, primarily because
attention was focused on the fossiliferous basal clay-pebble
conglomerate. However, these deposits do appear very clear-
ly in field photographs taken at that time. The layered de-
posits of hematite at the contact between Members A and B
of the Inapari Formation can be seen in Fig. 10b, as well as
a similarly prominent layer occurring at the contact between
the Holocene and Tertiary strata. A sheet-like layer of he-
matite was also observed by the senior author in a roadcut
leading to a bridge crossing the Rio Buyuyo, due south of
Porvenir and just east of the border with Peru (Fig. 8a). At
this site, the layer of hematite separates the silts of Member
B of the Inapari Formation from the underlying clays of
Member A, at the base of which occurs the characteristic
clay-pebble conglomerate. The concretionary zone of he-
matite that occurs in the highest soil profiles along the Rio
Beni also appears in the complete sections along the Rio Acre
(Fig. 10a). It may be of interest to note that the nodules of
hematite from this concretionary zone were being used as

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Campbell, Frailey, and Arellano L.: Rio Beni Geology 9

a

b

Figure 11. A comparison of the generalized geological sections along the a Rio Acre and b Rio Jurua. In b, the section at Pedra Preta, “a”
represents Tertiary clays; “b” is a “heavy conglomerate” from which came Tertiary fossils; “c” is yellow-greenish clays and silts; and “d”
represents buff fine sands and sandy silts. The contact between the different units was not clearly seen, according to the author, a, from
Campbell and Frailey, 1984; b, from Paula Couto, 1978. A series of specific sections for the Rio Jurua are presented in Simpson and Paula
Couto, 1981.

“gravel” to surface the road running south from Cobija (on
the Rio Acre) to Porvenir (on the Rio Tahuamanu) and the
Rio Buyuyo. ONERN (1977) states that iron concretions 1-
3 cm in thickness occur in the Inapari Formation (its Shirin-
gayoc Formation).

Campbell and Frailey (1984, in press) correlated the out-
crops exposed along the Rio Acre with those found along the
Rio Jurua in western Brazil (Paula Couto, 1978, 1983; Simp-
son and Paula Couto, 1981) (Fig. 1 lb). If that correlation is
correct and the deposits of the Rio Beni correlate with those
of the Rio Acre, then the deposits of the Rio Beni must also
correlate with those of the Rio Jurua. In that case, all of the
Quaternary deposits of the Monte Realm in Bolivia may be
referred to the Inapari Formation. The Inapari Formation,
and the Monte Depositional Realm, must then have a min-
imum areal coverage extending from its source area some-
where in the Peruvian Andes to the Rio Jurua to the north-
east, and southeasterly to the Brazilian Shield in eastern
Bolivia. Future fieldwork to demonstrate conclusively the
broad areal extent of the Inapari Formation is planned, but
for the moment this interpretation is proposed as a working
hypothesis.

The widespread deposits of the Monte Realm, i.e., the

Inapari Formation, were almost certainly deposited contem-
poraneously with those of the Sierra Realm. The stratigraphic
and lithologic similarities between the three members of the
two deposits also indicate that they must have been formed
under very similar circumstances. It is possible that the two
depositional realms merge into one in Peruvian territory to
the northwest, in which case the deposits of the Sierra Realm
will also be referable to the Inapari Formation. The long
distance separating the two depositional realms of the Rio
Beni, where riverbank outcrops are much lower than those
within the depositional realms, indicates that the deposits of
the lower Rio Beni could not have been derived from the
upper reaches of that river.

DRAINAGE SYSTEM ANALYSIS

It is possible to obtain a great deal of information regarding
the physiography of northern Bolivia through an analysis of
the present drainage system. For this we have used satellite
photographs (Figs. 2, 3), a map of the Departamento de La
Paz produced from satellite photographs (Instituto Geogra-
phico Militar, 1981), and the only satellite-based topographic
map presently available for any part of northern Bolivia (Rio

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Campbell, Frailey, and Arellano L.: Rio Beni Geology

Madre de Dios; Hoja SC 19-16, 1979, Institute Geographico
Militar, La Paz).

From these sources and field observations it is possible to
divide the course of the Rio Beni east of the Andes into four
regions (Fig. 2). Region I extends from Rurrenabaque through
the Sierra Realm, and in it the Rio Beni is initially charac-
terized as a braided river and subsequently as an incised,
meandering river. The braided portion is a result of the lateral
release of water after being channeled through the narrow
canyon cut through the front range. Where braided, the river
bottom is covered by large cobbles and boulders. J ust upriver
from Buena Vista the various channels coalesce into one and
continue as a single incised channel to a point just down-
stream from Altamarani, at the eastern limit of the Sierra
Realm.

Region II encompasses the river system downriver from
the end of Region I to approximately the Rio Madidi. In this
section the river is characterized as highly meandering, with
oxbow lakes, cutoff meanders, and meander scars occurring
almost everywhere on both sides of the main channel. The
course of the river in the lower half of this section is delimited
as a band of light gray cut through the surrounding dark gray
(Figs. 2, 3). This color difference is explained by the presence
of gallery forests growing on the natural levees formed by
the river. The growth of these forests is aided by the slightly
higher ground and better drainage afforded the terrain near
the main river channel. A similar effect is noted along the
Rio Madidi and Rio Biata, among others, that are also mean-
dering rivers of the highest degree (Figs. 1 , 2). The Rio Biata
can be seen to parallel the Rio Beni almost to the front range
of the Andes (Figs. 1-3), and evidence from satellite pho-
tographs suggests that its channel terminates in cutoff mean-
ders of the Rio Beni. During periods of flooding, the Rio
Biata must share the drainage of water coming through the
front range at Rurrenabaque. The Rio Beni may have become
the main river draining the region because of the additional
discharge it receives from the Rio Madidi, a factor that may
have allowed the Rio Beni to deepen its channel at a faster
rate than the Rio Biata, even though it may have had to cut
a longer channel through the Monte Realm.

Region III of the Rio Beni extends from approximately
the Rio Madidi to Ciudad California (Fig. 1, loc. 13). In this
region the river channel is incised, with long, straight stretch-
es alternating with broad, gentle curves. The beginning of
this section is just downriver from the southern limit of the
Monte Realm, and the river is actively downcutting through
the loosely consolidated Holocene sediments. It has not yet
reached that stage of development where extensive lateral
movements of the channel are possible; the channel is still
contained within a very narrow valley and there are very few
cutoff meanders or oxbow lakes.

In Region IV, from Ciudad California to Riberalta, the
Rio Beni is a meandering river, with abundant cutoff mean-
ders, oxbow lakes, and meander scars. In contrast to Region
II, with its natural levees built up above the surrounding
terrain, the meandering river in Region IV is constrained
within a narrow valley cut into the deposits of the Monte
Realm. The meanders are large and widely spaced initially,

but become smaller and more numerous approaching Ri-
beralta, as do the cutoff meanders and oxbow lakes. The
valley limits also become more clearly demarcated down-
stream. On satellite photographs the valley appears as a slightly
darker gray, narrow band cutting through the surrounding
light gray of the Monte Realm. A similar effect is perhaps
more clearly seen for the valley of the larger Rio Madre de
Dios just to the north (Figs. 2, 3), where prominent scalloping
of the valley walls clearly shows how the meandering river
is enlarging its valley through lateral erosion.

Outcrops in the Monte Realm occur wherever the river
channel curves into the constraining valley wall, undercutting
it and forming a cliff. In Region III the outcrops are widely
spaced, and all but the two northernmost occur on the eastern
side of the river. In Region IV the outcrops are more closely
spaced, and in many places alternate from one side of the
river to the other as the meanders are deflected from alternate
sides of the narrow river valley (Fig. 1 ). As would be expected,
the valley of the Rio Beni is broader and its channel more
meandering downstream from the confluence of the Rio Bia-
ta, although its development is clearly not equal to that of
the Rio Madre de Dios just to the north (Fig. 2).

After the juncture of the Rio Beni with the Rio Madre de
Dios, the enlarged river channel continues its meandering
course for a short distance, but gradually the meanders are
extended into long curves and then into a relatively straight
channel. This reflects the canyonization of the river as it
leaves the unconsolidated, fine-grained Holocene sediments
of the Amazonian lowlands and begins crossing the hard
crystalline rocks of the Brazilian Shield. All the rivers of
northern Bolivia drain into Brazil via the Rio Madre de Dios,
thus the rocks of the Brazilian Shield form the local base
level for the entire region.

THE HOLOCENE HISTORY OF

SOUTHWESTERN AMAZONIA
The available data are still too few to develop a definitive
Holocene history of southwestern Amazonia, but they are
sufficiently adequate for us to present our preliminary inter-
pretations. While recognizing that major questions remain
and that extensive fieldwork is still to be done, we feel that
a picture is emerging.

One of the primary objectives of the geological survey of
the Rio Beni was to determine if three cycles of Holocene
deposition and erosion were detectable in the alluvium de-
posited by the Rio Beni after it exited the front range of the
Andes and entered the essentially flat terrain of the eastern
lowlands, the region we have termed the Sierra Depositional
Realm. As we have described, three different lithologic units
do occur in the Sierra Realm, and they are similar in strati-
graphic sequence and lithologic composition to the three
members of the Inapari Formation occurring farther down-
stream in the Monte Depositional Realm and along the Rio
Acre. The three lithologic units of the two depositional realms
are at present correlative only temporally; the deposits are
not laterally contiguous anywhere in Bolivia (Fig. 3), and
fieldwork in Peru is necessary to resolve their relationship
to each other.

Contributions in Science, Number 364

Campbell, Frailey, and Arellano L.: Rio Beni Geology 11

Figure 12. Map of the altiplano of Bolivia and southern Peru. The
4000 m contour line delimits the altiplano, within which lie two
large lakes (Lake Titicaca and Lake Poopo) and two large dry salt
lakes (Salar de Coipasa and Salar de Uyuni). The hatched area in-
dicates elevations greater than 4000 m within and surrounding the
altiplano. The triangles mark the most likely position for an ice dam,
and the arrows indicate the direction of flow of the floodwaters out
of the altiplano.

As noted earlier, however, the area covered by alluvium
deposited by the Rio Beni within the Sierra Realm is very
small, less even than that immediately to the north and south
where alluvium has accumulated from only the eastern slopes
of the front range. To account for this lack of extensive al-
luvial deposits, we propose the occurrence of a catastrophic
flood that (1) removed preexisting alluvial deposits from the
area immediately east of the exit of the Rio Beni from the
front range, and (2) cleansed the principal intermontane val-

leys of the Rio Beni of loose alluvium, thereby reducing the
quantities of sediments available to form alluvial deposits
after the flood. In addition, we propose that this flood (3)
formed by scouring the series of depressions now occupied
by lakes (Figs. 2, 3) that occur in a straight line northeast of
Rurrenabaque, and (4) deposited the characteristic clay-peb-
ble conglomerate that occurs as a basal component of the
Holocene Inapari Formation at least as far north as Cruzeiro
do Sul, Brasil, on the Rio Jurua.

As the source of the floodwaters, we propose a late Pleis-
tocene lake sited in the Andean altiplano of Bolivia and Peru,
the antecedent of Lake Titicaca (Fig. 12). The Andean alti-
plano is a broad, almost flat region occupying an area of
almost 200,000 km2 between the western and eastern cor-
dilleras of the Andes. Elevations range from about 4000 m
at its northern and southern limits to a low point near it
center (Salar de Uyuni) of 3660 m. Lohmann (1970:754)
referred to the formation of the altiplano peneplain as an
“unexplained phenomenon,” suggesting that possibly post-
Pleistocene lacustrine and fluvial sedimentation redistrib-
uted Pleistocene glacial deposits to form the peneplain. He
also stated that the altiplano was once continuous across the
present La Paz valley, requiring that the erosion of the valley
of the Rio de La Paz be a postglacial event. He calculated
that approximately 20 km3 of Quaternary and older material
was removed from this valley during the past 10,000 year
period, or that material was removed at an annual erosion
rate of 2 x 106 m3/yr (Lohmann, 1970). The mean annual
precipitation for La Paz is 439 mm (Montes de Oca, 1983),
which is clearly insufficient to account for the erosion re-
quired to form the valley. The elevation of the divide sep-
arating the altiplano drainage from that of the Rio Beni, via
the Rio de La Paz, is only about 3970 m. The Altiplano has
no external drainage, and its only potential outlet would be
through the Rio de La Paz, if the water were high enough.

The evidence from our preliminary field observations sug-
gests that glacial ice coming out of the Cordillera Real (spe-
cifically, the Cordillera de La Paz where elevations reach over
7000 m) covered the eastern half of the altiplano. Reaching
a ridge south of La Paz, this glacial ice could have formed a
dam behind which collected glacial meltwaters. Alternative-
ly, a glacial ice cap may have formed over the central ridge
leading south from Lake Titicaca and the eastern altiplano,
meeting the glacial cap coming off the mountains somewhere
in the eastern altiplano. The occurrence of a widespread ice
cap would help explain the formation of the altiplano pe-
neplain, and topographic features such as numerous stream-
lined hills resembling rock drumlins suggest that in the east-
ern part of the altiplano there was ice flow toward the Lake
Titicaca basin. This hypothesized glacial ice cover is in con-
trast to current views (Clapperton, 1983) that hold that dur-
ing the last glaciation the mountain ice caps terminated in
separate piedmont lobes and did not cover the altiplano.

Till deposits documented in the valley of the Rio de La
Paz to below 3500 m (Dobrovolny, 1956, 1 962) indicate that
the upper reaches of this valley were filled at various times
during the Quaternary with glacial ice prior to the formation
of the altiplano peneplain (Lohmann, 1970). Clapperton

12 Contributions in Science, Number 364

Campbell, Frailey, and Arellano L.: Rio Beni Geology

(1979) gives a date of 3.27 myr for a till in the Rio de La
Paz valley that lies at an elevation of about 3900 m. Glaciers
in the Cordillera de La Paz now reach elevations as low as
4800 m on the northeast flank of the range, and 5300 m on
the southeast flank (Munoz Reyes, 1 977). The size differences
of the existing glaciers is a result of the large precipitation
differences that exist between the northeast and southeast
flank of the mountain range.

The Rio de La Paz is eroding headward into the altiplano
via its small headwater tributaries, principally the Rio Cho-
queyapu and Rio Achocalla. But the upper reaches of the La
Paz Valley do not assume the characteristic badland features
consistent with headward erosion in a semiarid climate.
Rather, the Rio Choqueyapu ends headward in large U-shaped
glacial valleys, whereas the Rio Achocalla is a smaller stream
that drains a huge bowl-shaped depression, the Achocalla
Valley, with cliff-like rims that in places pose walls with shear
drops of hundreds of meters. Except near presently existing
stream channels, the bottom of this depression, while gen-
erally moderately to steeply sloping, is not eroded into the
typical badland topography usually found in regions with
poorly indurated Quaternary sediments and semiarid cli-
mates. The Rio Choqueyapu and the Rio Achocalla are clear-
ly underfit streams (fide Dury, 1964), and we find that the
Achocalla Valley bears a striking resemblance to the plunge
pool of a giant waterfall (see Ahlfeld and Branisa, 1960: fig.
56).

Glacial lakes are known to have occurred in the altiplano,
and three have been named (e.g., Ahifield and Branisa, 1960;
Bowman, 1909; Lavenu, 1981; Montes de Oca, 1983; Ser-
vant, 1977; Servant and Fontes, 1978). The history of these
lakes, however, is not very clear. Lake Ballivian existed in
the northern part of the altiplano and is said to have been
50 percent larger than Lake Titicaca, thus covering an area
of 12,600 km2. It left deposits at an elevation of 3850 m. No
exact age for this lake has been determined. Lake Minchin
existed in the southern part of the altiplano and covered an
area of 60,000 km2 at an elevation of 3760 m. Its period of
maximum extension has been put at before 27,500 yr B.P.,
based on radiocarbon dates of calcareous deposits. Lake Tau-
ca covered an area of 43,000 km2 at an elevation of 3720 m
in the southern altiplano, and is said to have existed from
1 3,000 to 1 0,000 yr B.P. Lake Titicaca presently covers about
8400 km2 at an elevation of 3810 m in the northern part of
the altiplano. Clapperton (1983) and Lavenu et al. (1984)
report lake levels even higher than 3850 m, with the latter
describing newly discovered lake features at an elevation of
3950 m. This is 140 m above the present level of Lake Ti-
ticaca and only 20 m less than what would be required for
a lake to overflow the lip of the altiplano. Although each of
the glacial lakes is said to have existed at different intergla-
cials, is there a possibility that perhaps they represent instead
stillstands of a single great lake that covered all of the alti-
plano?

We propose that just such an extensive glacial lake did
form after 14,000 yr B.P. when glacial ice in the Andes re-
ceded rapidly (Emiliani et al., 1975; Mercer, 1977). Unless
blocked by glacial ice in the western altiplano, this glacial

lake may have extended southward to cover an area in excess
of 150,000 km2. As the lake grew in size it overflowed the
edge of the glacial ice in the region of La Paz, and the over-
flowing water began forming the large horseshoe-shaped ba-
sin with high, vertical sides which is the Achocalla Valley.
At some point the system broke down and a large portion
of the remaining lake waters drained catastrophically. Per-
haps this resulted from a 200 m uplift of the altiplano north-
east of La Paz cited by Lohmann (1970:754) as possibly being
contemporaneous in part with the Holocene erosion of the
valley of La Paz.

Once the ice dam was breached, the water would have
cascaded down the narrow intermontane valley of the Rio
de La Paz and into the Rio Beni, dropping from an elevation
of about 4000 m to less than 250 m in a distance of ap-
proximately 350 river-km. At several points in the eastern
cordilleras the Rio Beni flows through very narrow canyons
cut transversely through high ridges. These narrow canyons
would have restricted the rate of flow, and the water would
have formed temporary lakes filling numerous intermontane
valleys. The narrow canyon of the front range, which is only
about 200 m wide, is just such a constriction. Features visible
on satellite photographs, such as what appear to be high
plateaus with large, abandoned river channels and possibly
giant ripple marks downstream from an anticline composed
of Tertiary rocks, suggest that the large valley just west of
the front range (Fig. 2, with large white cloud in center) was
indeed filled with water. This flooded valley would have
maintained great hydraulic pressure on the floodwaters flow-
ing through the narrow canyon of the front range. And be-
cause the valley would have served as a giant reservoir, the
flow through this canyon onto the lowlands may have con-
tinued for some time.

The floodwaters would have emerged onto the flat low-
lands as a gigantic, high-pressure stream, much like water
passing through a sluice gate in a dam. Carrying away any
alluvial deposits in its path, the narrow stream of water scal-
loped out the chain of rectangular depressions now occupied
by lakes that so prominently form a straight line pointing
away from the canyon exit (Figs. 1-3). A well-known alter-
native explanation for this chain of lakes is based on pre-
sumed fractures of basement rocks (Montes de Oca, 1983:
152), but it is difficult to see how basement fracturing could
be expressed through a surficial cover of Recent alluvium in
an area that is regularly inundated by floodwaters and is
without recorded seismic activity. Even Laguna Rogaguado
(Fig. 3) and the lakes surrounding it, which lie to the northeast
of and are in line with the chain of rectangular lakes, may
have been formed by the erosive powers of this stream of
water.

On satellite photographs there is a strong hint of an alluvial
fan of standard shape with its center near the Rio Beni and
extending northeastward in a large semicircle. A large, dark
wedge-shaped slice is removed right of center. It should be
noted that although the present course of the main channel
of the Rio Beni is at an angle to the chain of lakes, the
structure of the canyon through the front range is such that
a high volume of water would be forced in the direction of

Contributions in Science, Number 364

Campbell, Frailey, and Arellano L.: Rio Beni Geology 13

Figure 13. a. The partial associated skeleton of a toxodont lies in
situ in Tertiary (late Miocene, Huayquerian) clays along the Rio
Acre. The two lower jaws, a scapula, and several leg bones are visible,
b. In the foreground, the clay-pebble conglomerate, or Acre Con-
glomerate Member of the Inapari Formation, is seen resting on Ter-
tiary clays at a locality on the Rio Acre. Channel deposits and strat-
ified clays of Member A of the Inapari Formation can be seen in the
background.

the lakes (Fig. 2). This course is followed today by one of
the braided river channels before it turns north-northeast-
ward and joins with the main river channel. After exiting the
front range, the floodwaters would have followed a general
northward path as a sheet flood, disrupting existing drainage
systems.

There is no reason to assume that the pre-Holocene Qua-
ternary drainage system of southwestern Amazonia bore any
resemblance to that of the area today. In fact, it may be
expected that the pre-Holocene drainage of northern Bolivia
flowed due north over the relatively soft Tertiary clays, skirt-
ing the western edge of the Brazilian Shield rather than cut-
ting a canyon through its hard crystalline rocks, much like
the Rios Itenez and Mamore do in part today. If such were
the case, the later deposition of the Inapari Formation created
a dam, closing off direct northern drainage from Bolivia and

forcing the rivers to exit Bolivian territory by crossing over
the Brazilian Shield, the course now followed by the Rio
Madre de Dios before it becomes the Rio Madeira and drains
into the Rio Amazonas. The Inapari Formation, extending
as a blanket over older strata from the Peruvian Andes to
the Brazilian Shield, may be effectively covering all evidence
of pre-Holocene Quaternary drainage systems.

Racing northward, the sheet flood hypothetically stripped
the soil from the surface, exposed the underlying unaltered
Tertiary deposits, and deposited the characteristic basal clay-
pebble conglomerate of the Inapari Formation. Prior to the
deposition of the Inapari Formation the general elevation of
the region would have been at least 20-35 m lower than
today and the terrain was probably even more level than at
present. Therefore, it can be assumed that the rivers were at
least as easily susceptible to flooding as they are at present,
and that any sheet flood could not possibly have been con-
tained in such shallow river valleys.

Of special importance is the probability that in its short-
lived passage the sheet flood deposited the characteristic
clay-pebble conglomerate with its reworked Tertiary fossil
vertebrates and wood that occurs only at the base of the
Holocene deposits. Simpson and Paula Couto (1981) and
Frailey (1980) remarked on the unusual conditions of the
fossil vertebrates in this basal Holocene deposit. Some of the
fossils are very water worn, indicating long distance trans-
port, but others cannot have been transported far, or re-
worked extensively. For example, Simpson and Paula Couto
(1981:19) state that “. . . glyptodont scutes may have been
worn almost formless but may also occur in large, unworn
clusters in which the plates have not been separated at su-
tures; . . . .” They also reported finding no associated fossil
skeletons. Campbell and Frailey (in press) and Frailey (1980)
reported on rich fossil localities on the Rio Acre where they
did find associated vertebrate skeletons, but these were in
situ in the Tertiary strata, not part of the clay-pebble con-
glomerate (Fig. 1 3a). They were, however, at the same phys-
ical level relative to the river channel as the clay-pebble
conglomerate (Fig. 1 3b).

We propose that the Tertiary fossils in the clay-pebble
conglomerate were reworked and redeposited near their point
of origin by the churning sheet flood, and that the reported
differences in wear of the fossils occurred at the time of their
primary deposition. In the case of the aforementioned glyp-
todont scutes, the well-worn, isolated scutes may have under-
gone long distance transport prior to their fossilization in the
Tertiary, while the partial carapaces composed of many scutes
were fossilized in the Teriary without being transported far,
if at all. Permineralized as a block, the partial carapace was
perhaps too strong to be broken up into individual scutes by
the passing floodwaters before they were redeposited a short
distance from their original site. But at the same time, ele-
ments of associated fossil skeletons present in the Tertiary
deposits would probably have been dispersed before being
redeposited in the clay-pebble conglomerate.

Campbell and Frailey (1984, in press) proposed a flooding
mechanism to remove the soil from the Tertiary strata and
deposit the clay-pebble conglomerate noted along the Rio

14 Contributions in Science, Number 364

Campbell, Frailey, and Arellano L.: Rio Beni Geology

Acre and Rio Jurua, but at the time they did not have a single
catastrophic flood in mind. Instead they proposed that gen-
eral massive flooding, sparked by rapid melting of Andean
glaciers at the end of the Pleistocene (Emiliani et ah, 1975)
and occurring over a short time period, produced the ob-
served geological features. A single catastrophic flood is a
more parsimonious hypothesis to explain the formation of
the clay-pebble conglomerate, as well as other geological fea-
tures in the region. It is necessary, however, to determine
where the coarse debris and large bedform features expected
from such a flood are located. It is probable that the evidence
necessary to document the flood will be found in the moun-
tains rather than in the forested and swampy lowlands where
Holocene deposits may be covering all but a few traces of
the erosional and depositional features of the flood. The search
for such evidence is in progress, but until it is available we
recognize the speculative nature of our proposal. Cata-
strophic flooding of enormous scale resulting from abrupt
draining of glacial lakes has been recorded elsewhere (Baker
andNummedal, 1978; Bretz, 1923; Bretz et ah, 1956;Malde,
1968), and large-scale catastrophic floods have even been
proposed to explain certain geological features on Mars (Bak-
er, 1978).

A hypothesis for the deposition of the three members of
the Inapari Formation was presented by Campbell and Frail-
ey (1984, in press). The hypothesis of a catastrophic flood
would modify their interpretation of events slightly in that
the fossiliferous basal clay-pebble conglomerate of Member
A of the Ihapari Formation is now seen as a unique deposit
formed at a specific time prior to the deposition of the re-
mainder of Member A. To recognize this distinction we pro-
pose that this unit be named the Acre Conglomerate Member
of the Inapari Formation, after the river where its distinc-
tiveness was first clarified. The type locality is on the south
bank of Rio Acre, 2 km east of the confluence of Rio de Los
Patos.

Following the passage of the sheet flood, sediments that
formed the Inapari Formation were washed eastward out of
the Peruvian Andes by the meltwaters of the disappearing
glaciers. More field data are required to pinpoint the exact
source of these sediments. Large-scale seasonal flooding last-
ing from about 10,000 yr B.P. to about 5000 yr B.P. was
proposed by Campbell and Frailey ( 1 984, in press) to account
for the highly channeled, variably unstratified and stratified
clay deposits of Member A of the Inapari Formation. This
proposal was based in part on the model developed by Kutz-
bach (1981) to explain early Holocene pluvials in Eurasia.

Rather specific conditions must have existed for the rel-
atively pure clays of Member A of the Inapari Formation to
be deposited over such a broad region. This probability is
increased by the fact that Member A in the Sierra Realm is
also a channeled clay deposit. It could be expected that the
sediment load of rivers entering the Amazonian lowlands
had a sizable clay fraction, a feature not unusual for waters
draining large areas of recently exposed glacial deposits in
high mountains. However, the deposition of clays requires
essentially still water. This indicates that inundated areas in
the region drained slowly, except in restricted zones where

the paleochannels of silts, sands, and clay-ball conglomerates
(Fig. 7) indicate fairly weak to strong current flow.

One possible cause of large-scale still water may have been
that the preexisting, i.e., pre-Holocene, drainage system was
blocked by sediments or otherwise disrupted by debris from
the catastrophic sheet flood as described above. This would
not be unexpected if, as suggested, the terrain were more
flattened than today. Taken together with the increased sea-
sonal precipitation and more rapid and extensive snowpack
melt proposed by Campbell and Frailey ( 1 984), an essentially
nonexistent drainage system throughout southwestern Ama-
zonia could have accentuated the development of broad,
anastomosing rivers with minimal gradients and extensive
swamplands susceptible to ready inundation. These condi-
tions would lead to the deposition of widespread clay de-
posits.

After a change in climate about 5000 yr B.P., and a sub-
sequent period of erosion, Members B and C of the Inapari
Formation were deposited. The lithologic similarity between
these two units indicate that they were deposited under sim-
ilar conditions, and their lithologic differences from Member
A, i.e., silts and fine sands instead of clays, indicate that these
conditions were different from those that prevailed earlier in
the Holocene. The break between the last two periods of
deposition was placed at about 2800 yr B.P., under the as-
sumption that the break in deposition represented a signif-
icant climatic change that could be expected to correlate with
Holocene climatic events recorded elsewhere (Denton and
Karlen, 1973; Fairbridge, 1976; Mercer, 1977; Wendland
and Bryson, 1974). The worldwide climatic anomalies ex-
perienced during 1983 (Philander, 1983a, b; Gill and Ras-
musson, 1983) clearly demonstrated how, if not why, such
correlations may occur.

The northward drainage of the eastern lowlands of Bolivia
must have been completely blocked by the time Member C
of the Inapari Formation was deposited, leaving much of the
region south of the Monte Realm waterlogged until drainage
was established over the Brazilian Shield. The drainage pat-
tern that has emerged since that time is one of a series of
major rivers, often flowing for long distances in roughly par-
allel courses (Figs. 1, 3), then converging in the northeastern
comer of the country and crossing the Brazilian Shield. Sec-
ondary drainage follows a classic dendritic pattern, indicating
uniformity of substrate and lack of structural control.

The imposition of the drainage system upon the Inapari
Formation can be seen rather clearly in the regions of the
Rio Beni described earlier. These regions represent different
stages of development, or maturity, of the river. Region III
is typical of a young river that is actively eroding downward,
while Region IV reflects a more mature stage, with a mean-
dering channel developing and occupying a broader valley.
A similar effect is reflected in the valley of the Rio Madre
de Dios, where, of that segment of the river course shown in
Fig. 3, the eastern half is far more meandering within a more
clearly developed valley than the western half.

The highly meandering nature of Region II of the Rio Beni
is a result of the Inapari Formation acting as a local base
level for the river, just as the crystalline rocks of the Brazilian

Contributions in Science, Number 364

Campbell, Frailey, and Arellano L.: Rio Beni Geology 15

Shield act as the local base level downriver from the conflu-
ence of the Rio Beni and Rio Madre de Dios. The natural
levees so prominent in Region II result from deposition of
sediment as the river overflowed its low banks during periods
of flooding and flowed laterally, unrestrained by valley walls.
Such flooding is probably accentuated by the fact that in
Region III the river enters into a narrow valley that can only
accommodate a portion of the river’s discharge during pe-
riods of flooding.

The river terraces noted along the Rio Beni within the
Monte Realm probably resulted from temporary local base
levels established and then eroded away in the course of the
river over the Brazilian Shield. That terraces of approxi-
mately similar height above the river can be found at many
localities along the river within the Monte Realm suggests
that some of the temporary local base levels lasted for a
significant period of time. As one bit of evidence for local
control over the formation of these terraces we cite the ab-
sence of a comparable series of terraces along the Rio Acre.
In fact, very few places along the Rio Acre upstream from
Cobija can be considered as river terraces.

IMPLICATIONS FOR OTHER RESEARCH

Clearly, more documentation of the proposed catastrophic
flood and further data regarding the integrity of the Inapari
Formation are required before any definitive statements can
be made about the Quaternary history of southwestern Ama-
zonia. Nevertheless, the geological data from the Rio Beni
reemphasizes the importance of studies of the Quaternary
geology of Amazonia to other fields of research as noted by
Campbell and Frailey (1984, in press). For example, one
important question we can raise is, just how old are the
tropical Amazonian forests as we know them? The traditional
view is that the forests are millions of years old (e.g., Abelson,
1983). The newly popular hypothesis of “tropical forest re-
fugia,” which holds that some areas of forest are very old,
having survived intact during the Pleistocene, while sur-
rounding areas of forest were converted to savannas by cli-
matic changes (see papers in Duellman, 1979, and Prance,
1982) is based almost entirely on the distributions of plants
and animals. Without supporting geological data, any inter-
pretations of the age of Amazonian ecosystems must be con-
sidered speculative.

As another example we may cite the use of the “tropical
forest refugia” hypothesis to determine the location of areas
of high priority or conservation (Lovejoy, 1982). The Rio
Acre region is proposed by some (e.g.. Brown, 1982) to have
been a tropical forest refugium during the Pleistocene, and
therefore remained as a stable forest community into the
present. But this proposal is untenable given the possibility
that the region was swept clear by a passing flood and because
the region is covered by the Holocene Inapari Formation,
the youngest deposits of which may be less than two thousand
years old. We agree that there is a good probability that
tropical forest refugia existed during the Pleistocene, and that

these areas should receive priority status regarding conser-
vation. But it is critical for these areas to be delimited first
on a geological, not a biological, basis.

In this regard, it is interesting to note that the greatest
avian species diversity for any locality in the world, over 500
species, is to be found in the Tambopata Reserve of Peru
(Parker, 1982), located just 210 km south of the Rio Acre
and 150 km southwest of the Rio Buyuyo site. Campbell and
Frailey (1984, in press) suggested that there is a good prob-
ability that this reserve, at an elevation of only 260 m, is
resting on surficial deposits of the Inapari Formation. If so,
the high diversity recorded there cannot be a result of long-
term environmental stability.

Similarly, studies of such disparate topics ranging from
Amazonian biogeography to pedology cannot move forward
without a better understanding of the geological history of
the region. Unfortunately, a review of a recent important
symposium on Amazonia (Donsereau, 1984) held no men-
tion of Quaternary geology, an indication of how the subject
is neglected. This can only be to the detriment of all fields
of research relating to Amazonia that are, or should be, de-
pendent on a thorough understanding of regional geological
history.

Even research into the paleoethnography of Amazonia is
incomplete without basic studies of late Quaternary geology.
Extensive flooding and depositional events in southwestern
Amazonia must have had a dramatic impact on Amerindians
in the region. These rather severe environmental changes
may explain the absence of early Holocene archaeological
sites in Amazonia (Meggers, 1979, 1982), and the fact that
few sites in Amazonia with terra preta soils are reliably dated
beyond about 2000 yr B.P. (Eden et al., 1 984). The geological
events proposed and described above may also be the basis
for some of the myths of Amerindians that speak of periods
of great floods (Meggers, 1982). One such myth, said to orig-
inate with the “Huni Kui” tribe of the Amahuaca Indians
that lived in southwestern Amazonia in the region between
the Rio Acre and Rio Jurua may even be an “eyewitness”
account of the catastrophic flood we have proposed:

In the dim and ancient past beyond recall, when man
could still talk with animals, our people had many vil-
lages and lived in peace with abundance of everything.
They lacked nothing and lived in happiness on the sandy
shore of the great river where the water meets the sky.

One day there came a great storm, worse than ever
before. It rained day and night. Everything stopped and
the people went to their houses. Thunder and lightning
came with a terrible wind, destroying the houses.

The sky broke and fell down. The earth went up into
the sky. Everything died except some crabs in a hole.
No other life remained. The land became sky; the sky,
land. Then the sky returned to its place and took the
spirits of the dead with it up into the sky. There the
spirits lived happily, but on the earth there was nothing
but a few crabs. [Lamb, 1974:120-121]

16 Contributions in Science, Number 364

Campbell, Frailey, and Arellano L.: Rio Beni Geology

SUMMARY

A geological survey along the Rio Beni in northern Bolivia
has demonstrated the existence of Quaternary sediments that
were deposited in two widely separated depositional realms:
the Sierra Realm, with sediments derived from the Bolivian
Andes; and the Monte Realm, with sediments derived from
the Peruvian Andes. The deposits in the two depositional
realms were probably deposited contemporaneously, and the
same sequence of deposits occurs in both: a lower unit of
channeled, variably stratified clays and two upper units of
blocky silts and fine sands.

The sediments of the Monte Realm are referred to the
Inapari Formation of ONERN (1977), a series of alluvial
deposits assigned to the Holocene by Campbell and Frailey
(1984, in press). The channeled, variably stratified clays of
Member A of the Inapari Formation were deposited from
about 10,000 yr B.P. to about 5000 yr B.P. Following a period
of erosion, and a change in the environment of deposition,
the blocky silts and fine sands of Members B and C of the
Inapari Formation were deposited. A disconformity sepa-
rating these two members represents a period of erosion, the
age of which is put at about 2800 yr B.P. The Inapari For-
mation appears to cover at least that part of southwestern
Amazonia extending from the Peruvian Andes northeast to
the Rio Jurua (to Cruzeiro do Sul, Brasil) and southeast to
the Brazilian Shield.

The deposition of the Inapari Formation formed a dam
across northern Bolivia, forcing the regional drainage north-
eastward, over the crystalline rocks of the Brazilian Shield.
The imposition of the present drainage system upon the Ina-
pari Formation is reflected in the varying stages of devel-
opment of the river valleys in northern Bolivia, with the
course of the Rio Beni east of the Andes being divisible into
four clearly distinct regions.

A catastrophic flood resulting from the sudden draining of
glacial Lake Titicaca is proposed to account for (1) the lack
of alluvial deposits extending eastward from the front range
of the Andes at the point where the Rio Beni enters the flat
lowlands; (2) the prominent chain of lakes extending north-
eastward in a straight line from the opening of the Rio Beni
onto the flat lowlands; and (3) the deposition of the char-
acteristic clay-pebble conglomerate with Tertiary fossil ver-
tebrates and wood, herein named the Acre Conglomerate
Member of the Inapari Formation.

The documentation of Holocene deposits over wide re-
gions of southwestern Amazonia has broad implications for
numerous fields of research, including such disparate fields
as biogeography, conservation, ecology, hydrology, pedology,
and paleoethnography.

ACKNOWLEDGMENTS

Fieldwork for this research was supported by the National
Science Foundation (DEB 78-03122 and DEB 78-05861),
the National Geographic Society (Grants Nos. 1776 and 2002),

and the Servicio Geologico de Bolivia (GEOBOL). We thank
Lidia Lustig and Alfonso Medina for their assistance in the
field, and Raul Carrasco C. and GEOBOL for encouraging
our fieldwork and making it possible. We thank Carlos Son-
nenschein of Riberalta for his many courtesies and his con-
tributions to our work.

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fauna: Its origin, evolution, and dispersal. Monograph
no. 7, 485 pp. University of Kansas, Museum of Natural
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Campbell, Frailey, and Arellano L.: Rio Beni Geology 17

Eden, M., W. Bray, L. Herrera, and C. McEwan. 1984.
Terra preta soils and their archaeological context in the
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Emiliani, C., S. Gartner, B. Lidz, K. Eldridge, D. Elvey, T.
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Fairbridge, R.W. 1976. Shellfish-eating Preceramic Indians
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mals from the Cenozoic of Acre, Brazil III — Pleistocene
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episodes of the Holocene. Quaternary Research 4:9-24.

Accepted for publication 8 January 1985.

18 Contributions in Science, Number 364

Campbell, Frailey, and Arellano L.: Rio Beni Geology

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CONTRIBUTIONS IN SCIENCE

VASCULAR PLANTS OF THE CHANNEL ISLANDS, OF SOUTHERN

CALIFORNIA AND GUADALUPE ISLAND,

BAJA CALIFORNIA, MEXICO

aJIasaflSS

.■ ■ •:..- . j‘;h; ■ .; v ■•?► -

,: • fepj, p?tij

-r- ®!lMPTpLsdi«

SERIAL PUBLICATIONS OF THE
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

The scientific publications of the Natural History Museum of Los Angeles County have been
issued at irregular intervals in three major series; the articles in each series are numbered
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® Contributions in Science, a miscellaneous series of technical papers describing orig-
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Gary D. Wallace
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Printed at Allen Press, Inc.. Lawrence. Kansas

VASCULAR PLANTS OF THE CHANNEL ISLANDS OF SOUTHERN
CALIFORNIA AND GUADALUPE ISLAND,

BAJA CALIFORNIA, MEXICO

Gary D. Wallace

Contributions in Science, Number 365
Natural History Museum of Los Angeles County
13 June 1985

ISSN 0459-8113

Natural History Museum of Los Angeles County
900 Exposition Boulevard
Los Angeles, California 90007

CONTENTS

ABSTRACT 1

INTRODUCTION 1

THE VASCULAR PLANTS OF THE CHANNEL ISLANDS OF
SOUTHERN CALIFORNIA AND GUADALUPE ISLAND,

BAJA CALIFORNIA, MEXICO 2

Vascular Cryptogams 3

Gymnosperms 4

Dicotyledons 4

Monocotyledons 29

FLORISTIC RELATIONSHIPS 34

APPENDIXES 40

Appendix I. Selected Exsiccatae and Herbaria Housing Insular

Collections 40

Vascular Cryptogams 41

Gymnosperms 42

Dicotyledons 42

Monocotyledons 83

Appendix II. Index to the Disposition of Synonyms, Misidentifi-

cations, and Taxa Incertae sedis 91

ACKNOWLEDGMENTS 133

LITERATURE CITED 133

MAPS AND TABLES

Map 1. Channel Islands of Southern California and Isla Guadalupe,

Mexico 38

Table 1. Features of the Channel Islands of Southern California

and Isla Guadalupe, Mexico and Floristic Data for Each 39

Table 2. Shared Native and Endemic Taxa Between Island Pairs 40

VASCULAR PLANTS OF THE CHANNEL ISLANDS OF SOUTHERN
CALIFORNIA AND GUADALUPE ISLAND,

BAJA CALIFORNIA, MEXICO

Gary D. Wallace1

ABSTRACT. Vascular plant taxa of the Channel Islands of southern
California and Guadalupe Island, Mexico, are treated, to include
their distribution among the islands, citation of verified specimens
from several herbaria, and a detailed account of earlier synonymous
taxa, misidentified specimens, and taxa incertae sedis based on her-
barium specimens and literature citations. A total of 621 native
vascular taxa include 137 endemics and 227 introduced taxa.

RESUMEN. Se tratan los taxa de plantas vasculares de las Channel
Islands de California Sur y de la Isla Guadalupe, Mexico, incluyendo
sus distribuciones dentro de las islas, citacion verificado de especi-
menes de diversos herbarios, y una cuenta detallada de sinonimos
previos, especimenes mal identificados, y taxa incertae sedis basada
en especimenes de herbarios y citas de la literatura. Se reporta un
total de 621 taxa vasculares indigenos, incluyendo 137 endemicos,
y 227 introducidos.

INTRODUCTION

There have been numerous papers over the past century
dealing with the plants of the Channel Islands. These were
often the results of single visits to a specific island (e.g.,
Greene, 1887a), summary works primarily based on previ-
ously published works (e.g., Brandegee, 1890b; Eastwood,
1941), or works combining these approaches (e.g., Raven,
1963; Thome, 1967; Foreman, 1 967; Philbrick, 1972; Smith,
1976).

The islands treated here and a reference to a treatment of
the flora of each are as follows: San Miguel Island (Smith,
1976); Santa Rosa Island (Smith, 1976); Santa Cruz Island
(Smith, 1976); Anacapa Islands (Smith, 1976); San Nicolas
Island (Foreman, 1967); Santa Barbara Island (Philbrick,
1972); Santa Catalina Island (Thorne, 1967); San Clemente
Island (Raven, 1963); Isla Guadalupe (Eastwood, 1929). Each
of these deals, almost exclusively, with the flora of a single
island or group of islands but none has treated the flora of
all of the islands. Foreman (1967), Philbrick (1972), Thome
( 1 967), Raven ( 1 963), and Eastwood ( 1 929) were among those
authors to cite verifiable specimens. Difficulties have arisen
over the years in locating specimens upon which earlier and

Contributions in Science, Number 365, pp. 1-136
Natural History Museum of Los Angeles County, 1985

erroneous reports were based, in some cases these have not
yet been located. Raven ( 1 963) and Thome (1967) dismissed
these unsubstantiated reports. Philbrick (1972) consistently
documented each case of error and misidentification. There
was no question as to which reports Philbrick had seen. Phil-
brick (1972) documented the disposition of persistent errors
in the literature. Finally Philbrick (1972) consistently pro-
vided citation of specimens which could be available for the
monographic and floristic studies of others.

The most recent treatment of the flora of the Channel
Islands as a whole was that done by Eastwood (1941). This
work was based on herbarium specimens, published reports
of collections, and apparently some lists compiled by bota-
nists of specimens they had collected. Eastwood (1941) cited
no collections, making confirmation of her insular records
difficult. Apparent errors were perpetuated by this work and
original determinations were further obscured by the fact that
Eastwood occasionally submerged or changed these with no
annotation of the specimens she saw. Because of some errors
in Eastwood (1941), additional inconsistencies may have been
arbitrarily attributed to her list. Some errors attributed to
Eastwood’s list are undoubtedly the result of her inclusion
of data from earlier published and unpublished works. Among
major floristic works, Munz and Keck ( 1 959) only occasion-
ally noted specific insular distributions of taxa. Munz ( 1974)
in addition to this was inconsistent in including insular taxa
from the northern islands. Smith (1976) gave an effective
treatment of the plants of the northern group of islands in
his regional flora.

The Channel Islands are an important phytogeographical
unit of California. Many of the insular taxa are in need of
systematic and evolutionary study as indicated by Raven
(1963), Thome (1967), Philbrick (1972), and Smith (1976).
Several taxa reach their northern or southern range limits

1. Botany Section, Natural History Museum of Los Angeles
County, 900 Exposition Blvd., Los Angeles, California 90007.

ISSN 0459-8113

among the Channel Islands and Guadalupe Island, Mexico.
The outlier populations may provide important data to assess
patterns of variation and evolution. Treatments by Davis
(1980) and Philbrick (1980) depict some of the intricacies of
evolutionary development found among insular taxa.
Knowledge of the occurrence and distribution of the insular
plant taxa, as well as the availability of specimens, is vital
to workers in these areas.

The high incidence of endemism is apparent in several
genera, most notably Eriogonum (Polygonaceae), Dudleya
(Crassulaceae), Arctostaphylos (Ericaceae), Galium (Rubi-
aceae), Malacothrix (Asteraceae), Hemizonia (Asteraceae),
Phacelia (Hydrophyllaceae), and Lotus (Fabaceae). Discus-
sions of the significance, origins, and distributions of the
endemic plants on the Channel Islands may be found in
Stebbins and Major (1965), Raven (1967, 1977), and Phil-
brick (1980). General information about the vegetation of
the islands may be found in Philbrick and Haller (1977), as
well as in the several treatments for specific islands. Westman
(1983) discussed the structure of the insular xeric shrublands
and compared them to similar mainland sites. Philbrick (1967)
and Power (1980) provide introductions and sources for those
interested in the diverse aspects of geological, natural and
evolutionary history of the Channel Islands.

The Natural History Museum of Los Angeles County has
had a long association with the study of the Channel Islands.
The combined collections of AHFH, USC, and LAM at LAM
contain significant numbers of insular collections of M.B.
Dunkle, F.R. Fosberg, and F.H. Elmore as well as numerous
specimens collected by R. Moran, B. Trask, I.W. Clokey, and
P.J. Rempel among others (herbarium acronyms throughout
are as given in Holmgren et al., 1981). One of the most active
centers for floristic treatment of the Channel Islands is cur-
rently the Santa Barbara Botanic Garden. Dr. Ralph Phil-
brick and his co-workers have made extensive and intensive
collections of the northern islands as well as Santa Barbara
Island. Rancho Santa Ana Botanic Garden has been the pri-
mary center for the floristic study of Santa Catalina and San
Clemente Islands. Dr. Robert F. Thome has made exhaustive
collections on Santa Catalina Island and has made several
collection trips to the other islands. The current study of
Guadalupe Island, Mexico, is almost exclusively centered at
the San Diego Museum of Natural History. Dr. Reid Moran
made numerous trips over the years to the island and is the
recognized authority on the flora of Guadalupe. Several other
institutions have significant insular collections which sup-
ported earlier research specialties or interests (AHFH, CAS,
CM, DS, F, GH, MO, ND-G, NY, US). The Natural History
Museum and the Santa Barbara Museum contain both his-
torical and current specimens, a reflection of sustained in-
terest in the islands.

The objectives of this paper are several. One is to provide
a current guide to the floras of each of the islands of this
phytogeographically important region. This is done in a man-
ner which clearly distinguishes between those records based
upon available herbarium specimens and those based on
reports from the literature. Another goal is to present a pic-
ture of the floristic relationships among the islands. Finally

I wanted to draw the attention of systematists to the insular
distributions of native taxa and to the numerous misiden-
tifications and synonymous treatments of those taxa. Their
aid in the verification and correction of data in the two ap-
pendixes will greatly clarify our knowledge of the insular
floras.

This treatment is primarily based on verified herbarium
specimens. Specimens cited are deposited in one or more of
the following collections: CAS-DS, CM, F, GH, JEPS, LAM
(includes AHFH and USC), MO, ND-G, NY, RSA-POM,
SBBG, SBM, UC, UCSB, UCSB-SCIR, US, and the Pacific
Missile Test Center at Point Mugu, California.

Numerous monographic and floristic publications provid-
ed reports of insular plants. These publications are cited in
the table of vascular plants only when herbarium specimens
could not be found which verified the individual report, or
when the report could not be determined to be an error.

Personal communications of the occurrence of taxa on the
islands are not included nor are published sightings lacking
voucher specimens, reports from unpublished lists, or col-
lections not entered in an herbarium and available for loan.

In most cases, current monographic treatments served as
nomenclatural guides. Not all taxonomic judgments made
here will meet with approval. Comments including reference
to additional monographic treatments will be appreciated. I
hope monographers and curators will attempt to verify or
deny records known here from literature references citing the
specimens upon which the records are based. Appendixes I
and II are intended to facilitate this endeavor.

The paper is divided into three major sections, the table
of vascular plants, Floristic Relationships, and the appen-
dixes. Floristic Relationships contains Map 1, which shows
the spatial relationships among the islands, and two tables.
Table 1 lists some features of the islands treated here and
gives floristic summaries for each island. Table 2 indicates
the numbers of shared endemics and total number of shared
native taxa among the islands.

THE VASCULAR PLANTS OF THE CHANNEL
ISLANDS OF SOUTHERN CALIFORNIA AND
GUADALUPE ISLAND, BAJA
CALIFORNIA, MEXICO

The order of inclusion of the major plant groupings is Vas-
cular Cryptogams, followed by the Gymnosperms, Dicoty-
ledons, and Monocotyledons. Families, genera, and species
are alphabetically arranged within these groupings. Through-
out this paper, the abbreviations of authors’ names follow
those given by Munz and Keck (1959). In the first column
to the right of the name of the taxon there will be one of
three abbreviations: NA = native to the islands noted, EN =
native and more specifically endemic to islands noted, or
IN = introduced. Occasionally a taxon is endemic to the is-
lands treated here and one or a few other of the Mexican
islands; in such cases it will be noted as endemic and the
additional islands on which the species occurs are given in
parentheses after the name of the taxon. The next nine col-
umns correspond to one of each of the islands. A notation
in a column opposite the name of a taxon indicates a record

2 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

of occurrence based on herbarium material. The following
abbreviations correspond to the islands to avoid confusion
and occur throughout this paper: MI = San Miguel Island,
RO = Santa Rosa Island, CR = Santa Cruz Island, AN =
Anacapa Islands, NI = San Nicolas Island, BA = Santa Bar-
bara Island, CA = Santa Catalina Island, CL = San Clemente
Island, GU = Isla Guadalupe or Guadalupe Island Citation
of specimens and their locations for these records or only

location of specimens is noted in Appendix I. Occasionally
a number appears in the column; this corresponds to a nu-
merically indexed entry in the Literature Cited section. Most
of these reports are neither accepted nor denied but rather
are offered for the comment and study of others. Undoubt-
edly, many of these may be dismissed as the specimens upon
which they are based are located and identified or likely
herbarium sources are exhausted.

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur- Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence Island

Island

Island

Island

Island

Island

Island

Island

lupe

Vascular Cryptogams

Selaginellaceae

Selaginella bigelovii Underw.

NA

RO

CR

AN

CA

CL

Equisetaceae

Equisetum hyemale L. ssp. affine (En-

gelm.) Calder & Taylor

NA

CR

Equisetum laevigatum A. Br.
Equisetum telmateia Ehrh. var.

NA

RO

CR

CA

braunii Milde

NA

CR

CA

Aspidiaceae

Athyrium felix-femina (L.) Roth var.

sitchensis Rupr.

NA

31

CR

Cystopteris fragilis (L.) Bemh.

NA

CR

Dryopteris arguta (Kaulf.) Watt.
Polystichum munitum (Kaulf.) Presl.

NA

RO

CR

AN

CA

CL

ssp. munitum

NA

RO

CR

Polystichum munitum (Kaulf.) Presl

ssp. solitarium Maxon

EN

GU

Blechnaceae

Woodwardia fimbriata Sm. in Rees

NA

RO

CR

Polypodiaceae

Polypodium californicum Kaulf.

NA

RO

CR

AN

BA CA

CL

GU

Polypodium scouleri Hook. & Grev.

NA

CR

GU

Pteridaceae

Adiantum capillus-veneris L.

NA

RO

CR

AN

CA

Adiantum jordani C. Muell.
Adiantum pedatum L. ssp. aleuticum

NA

RO

CR

AN

CA

CL

(Rupr.) Calder & Taylor

Aspidotis californica (Hook.) Nutt, ex

NA

CR

Copel.

NA

CR

CA

Cheilanlhes clevelandii D.C. Eat.
Cheilanthes newberryi (D.C. Eat.)

NA

RO

CR

Domin

NA

CL

GU

Notholaena californica D.C. Eat.

NA

41

CA

GU

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 3

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Islam

Island

Island

Island

Island

Island

Island

lupe

Pellaea andromedaefolia (Kaulf.) Fee
var. pubescens D.C. Eat.

Pellaea mucronata (D.C. Eat.) D.C.

NA

RO

CR

AN

CA

CL

Eat. ssp. mucronata

NA

RO

CR

26

CA

GU

Pityrogramma triangularis (Kaulf.)

Maxon var. triangularis
Pityrogramma triangularis (Kaulf.)

NA

RO

CR

AN

CA

CL

GU

Maxon var. viscosa (D.C. Eat.)
Weath.

NA

RO

CR

NI

CA

CL

Pteridium aquilinum (L.) Kuhn var.

pubescens Underw.

NA

RO

CR

CA

Salviniaceae

Azolla filiculoides Lam.

NA

CA

Gymnosperms

Cupressaceae

Cupressus guadalupensis Wats. ssp.
guadalupensis

Cupressus macrocarpa Hartw. ex

EN

GU

Gord.

IN

CR

AN

CA

Juniperus californica Carr.

NA

GU

Pinaceae

Pinus muricata D. Don

NA

81

CR

Pinus radiata D. Don var. binata (En-

gelm. in Wats.) Lemmon

EN

GU

Pinus remorata Mason

NA

RO

CR

Pinus torreyana Parry ex Carr.

NA

RO

Dicotyledons

Aceraceae

Acer macrophyllum Pursh

NA

CR

Aizoaceae

Aptenia cordifolia (L. f.) N.E. Br.
Carpobrotus aequilaterus (Haw.) N.E.

IN

AN

CA

Br.

NA

MI

RO

CR

AN

NI

CL

Carpobrotus edulis (L.) Bolus

IN

NI

Malephora crocea (Jacq.) Schwant

IN

AN

Mesembryanthemum crystallinum L.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Mesembryanthemum nodijlorum L.
Tetragonia tetragonioides (Pall.)

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Kuntze

IN

MI

RO

CR

81

Amaranthaceae

Amaranthus albus L.

IN

31

RO

CR

CA

Amaranthus blitoides Wats.

IN

31

RO

CR

Amaranthus deflexus L.

IN

CR

Anacardiaceae

Lithraea molloides (Kell.) Engler

IN

CA

4 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Malosma laurina (Nutt, in T. & G.)

Nutt, ex Abrams

NA

CA

CL

GU

Rhus integrifolia (Nutt.) Benth. &

Hook.

NA

MI

RO

CR

AN

CA

CL

GU

Rhus ovata Wats.

NA

CR

CA

Schinus molle L.

Toxicodendron radicans (L.) Kuntze

IN

CR

CA

ssp. diversilobum (T. & G.) Thome

NA

MI

RO

CR

AN

CA

CL

Apocynaceae

Asclepias fascicularis Dene, in A. DC.
Sarcostemma cynanchoides Dene. ssp.

NA

CR

CA

hartwegii (Vail) R. Holm

NA

CA

Vine a major L.

IN

CR

CA

Araliaceae

Apiastrum angustifolium Nutt, in

T. & G.

NA

MI

RO

CR

AN

NI

CA

CL

Apium graveolens L.

IN

RO

CR

NI

CL

Berula erecta (Huds.) Cov.

NA

MI

41

26

Bowlesia incana R. & P.

IN

RO

CR

CA

CL

Caucalis microcarpa H. & A.

NA

CR

CA

CL

Conium maculatum L.

IN

CR

NI

31

Daucus pusillus Michx.

NA

MI

RO

CR

AN

NI

BA

CA

CL

GU

Foeniculum vulgare Mill.
Lomatium caruifolium (H. & A.)

IN

RO

CR

NI

CA

CL

Coult. & Rose

NA

MI

RO

CR

Lomatium insulare (Eastw.) Munz
Lomatium utriculatum (Nutt.) Coult.

EN

NI

CL

GU

& Rose

NA

RO

CR

Sanicula arguta Greene ex Coult. &

Rose

NA

MI

RO

CR

AN

NI

CA

CL

Sanicula crassicaulis Poepp. ex DC.

var. crassicaulis

NA

CR

CA

93

Sanicula hoffmannii (Munz) Bell

NA

MI

RO

CR

NI

Torilis nodosa (L.) Gaertn.

IN

MI

RO

CR

CA

Asteraceae

Achillea millefolium L.

NA

MI

RO

CR

AN

NI

BA

CA

CL

Achyrachaena mollis Schauer
Agoseris apargioides (Less.) Greene

NA

RO

CR

CL

ssp. apargioides

NA

RO

Agoseris grandiflora (Nutt.) Greene

NA

MI

RO

CR

Agoseris heterophylla (Nutt.) Greene

NA

RO

CR

GU

Amblyopappus pusillus H. & A.

NA

MI

RO

CR

AN

NI

BA

CA

CL

GU

Ambrosia acanthicarpa Hook.

IN

CA

Ambrosia camphor at a (Greene) Payne

NA

GU

Ambrosia chamissonis (Less.) Greene
Ambrosia psilostachya DC. var. call-

NA

MI

RO

CR

AN

NI

CA

CL

fornica (Rydb.) Blake in Tidestr.

NA

41

CA

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 5

Taxon

Occur-

rence

San

Miguel

Island

Santa

Rosa

Island

Santa

Cruz

Island

Anacapa

Island

San

Nicolas

Island

Santa

Barbara

Island

Santa San

Catalina Clemente
Island Island

Isla

Guada-

lupe

Anthemis cotula L.

IN

5

CR

CA

Artemisia californica Less.

NA

MI

RO

CR

AN

CA

CL

GU

Artemisia douglasiana Bess, in Hook.

NA

RO

CR

CA

Artemisia dracunculus L.

NA

CA

Artemisia nesiotica Raven

EN

NI

BA

CL

Aster chilensis Nees var. chilensis

NA

RO

CR

Aster exilis Ell.

NA

CR

66

Aster radulinus Gray

NA

RO

CR

Baccharis douglasii DC.

NA

MI

RO

CR

26

CA

Baccharis emoryi Gray

NA

CA

Baccharis glutinosa Pers.

NA

RO

CR

AN

CA

CL

Baccharis pilularis DC. ssp. consan-

guinea (DC.) C.B. Wolf

NA

MI

RO

CR

AN

NI

BA

CA

CL

Baccharis plummerae Gray

NA

CR

Baeriopsis guada/upensis J.T. Howell

EN

GU

Blennosperma nanum (Hook.) Blake

var. nanum

NA

CR

Brickellia californica (T. & G.) Gray

NA

CR

AN

CA

Centaurea cineraria L.

IN

CA

Centaurea melitensis L.

IN

MI

RO

CR

NI

BA

CA

CL

GU

Centaurea solstitialis L.

IN

31

CR

CA

Chaenactis glabriuscula DC. var. lanosa

(DC.) Hall

NA

RO

Chrysanthemum coronarium L.

IN

CR

NI

CA

Chrysanthemum frutescens L.

IN

CA

Cichorium intybus L.

IN

CR

Cirsium brevistylum Cronq.

NA

CR

Cirsium californicum Gray

NA

31

RO

CR

CA

Cirsium occidentale (Nutt.) Jeps.

NA

MI

RO

CR

NI

CA

CL

Cirsium ochrocentrum Gray

IN

CA

Cirsium proteanum J.T. Howell

NA

RO

Cirsium vulgar e (Savi) Ten.

IN

CA

Cnicus benedictus L.

IN

81

Conyza bonariensis (L.) Cronq.

IN

81

CR

NI

CA

CL

Conyza canadensis (L.) Cronq.

IN

MI

RO

CR

NI

CA

CL

Conyza coulleri Gray

NA

31

CR

CA

CL

Coreopsis gigantea (Kell.) Hall

NA

MI

RO

CR

AN

NI

BA

CA

GU

Corethrogyne filaginifolia (H. & A.)

ssp. filaginifolia

NA

MI

RO

CR

AN

NI

CA

Cotula australis (Sieber ex Spreng.)

Hook. f.

IN

RO

CR

CA

Cotula coronopifolia L.

IN

MI

RO

CR

NI

CA

Cynara scolymus L.

IN

CR

Encelia californica Nutt.

NA

CR

AN

CA

CL

Erigeron foliosus Nutt, var . foliosus

NA

MI

RO

CR

AN

CA

Erigeron foliosus Nutt. var. stenophyl-

lus (Nutt.) Gray

NA

MI

31

CR

AN

Erigeron glaucus Ker

NA

MI

RO

CR

AN

Erigeron sanctarum Wats.

NA

RO

26

6 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Taxon

Occur-

rence

San

Miguel

Island

Santa

Rosa

Island

Santa

Cruz

Island

Anaeapa

Island

San

Nicolas

Island

Santa

Barbara

Island

Santa San

Catalina Clemente
Island Island

Isla

Guada-

lupe

Eriophyllum confertijlorum (DC.)

Gray var. confertijlorum

NA

Ml

RO

CR

AN

CA

CL

Eriophyllum lanatum (Pursh) Forbes

var. grandijlorum (Gray) Jeps.

NA

GU

Eriophyllum nevinii Gray

EN

BA

CA

CL

Eriophyllum staechadifolium Lag. var.

arternisiaefolium (Less.) Macbr.

NA

MI

RO

CR

AN

Eriophyllum staechadifolium Lag. var.

depressum Greene

NA

MI

RO

CR

AN

Evax sparsiflora (Gray) Jeps.

NA

RO

31

CL

Filago arizonica Gray

NA

66

7

CA

CL

GU

Filago californica Nutt.

NA

MI

RO

CR

AN

CA

CL

GU

Filago gallica L.

IN

CR

CA

Gnaphalium beneolens A. Davids.

NA

MI

RO

CR

NI

CA

CL

Gnaphalium bicolor Bioletti

NA

43

RO

CR

AN

NI

CA

CL

GU

Gnaphalium californicum DC.

NA

RO

CR

AN

CA

Gnaphalium chilense Spreng.

NA

MI

RO

CR

AN

NI

CA

GU

Gnaphalium luteo-album L.

IN

MI

RO

CR

NI

CA

CL

Gnaphalium microcephalum Nutt.

NA

RO

CR

AN

CA

Gnaphalium palustre Nutt.

NA

CR

CA

CL

Gnaphalium purpureum L.

NA

RO

CR

81

CA

Gnaphalium ramosissimum Nutt.

NA

CR

Grindelia latifolia Kell.

NA

42

RO

98

AN

Grindelia robusta Nutt. var. robusta

NA

CR

NI

63

Grindelia stricta DC. ssp. venulosa

(Jeps.) Keck

NA

RO

31

Haplopappus canus (Gray) Blake

EN

CL

GU

Haplopappus detonsus (Greene) Raven

EN

RO

CR

AN

Haplopappus ericoides (Less.) H. & A.

ssp. ericoides

NA

42

Haplopappus palmeri Gray ssp. pachy-

lepis Hail

NA

CA

Haplopappus squarrosus H. & A. ssp.

grindelioides (DC.) Keck

NA

MI

RO

CR

CA

Haplopappus venetus (HBK.) Blake

ssp. furfuraceus (Greene) Hall

NA

NI

CA

CL

Haplopappus venetus (HBK.) Blake

ssp. sedoides (Greene) Munz

NA

MI

RO

CR

AN

Haplopappus venetus (HBK.) Blake

ssp. vernonioides (Nutt.) Hall

NA

MI

RO

CR

AN

NI

CA

CL

Helenium puberulum DC.

NA

CA

Helianthus annum L. ssp. lenticularis

(Dougl.) Ckll.

NA

CR

CA

Hemizonia Clementina Bdg.

EN

AN

NI

BA

CA

CL

Hemizonia fasciculata (DC.) T. & G.

NA

42

RO

CR

AN

BA

CA

CL

Hemizonia fitchii Gray

IN

CR

Hemizonia frutescens Gray

EN

GU

Hemizonia greeneana Rose ssp.

greeneana

EN

GU

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 7

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Hemizonia increscens (Hall ex Keck)

Tanowitz ssp. increscens

NA

RO

CR

Hemizonia palmeri Rose

EN

GU

Heterotheca grandiflora Nutt.
Hieraceum argutum Nutt. ssp. argu-

NA

CR

NI

CA

CL

turn

NA

RO

CR

Hypochoeris glabra L.

IN

RO

CR

CA

CL

GU

Jaumea carnosa (Less.) Gray

NA

MI

RO

CR

CA

Lactuca serriola L.

IN

RO

CR

NI

CA

Laslhenia californica DC. ex Lindl.

NA

MI

RO

CR

AN

NI

BA

CA

CL

GU

Lasthenia coronaria (Nutt.) Omduff
Lasthenia glabrata Lindl. ssp. coulteri

NA

GU

(Gray) Omduff

NA

RO

31

Layia glandulosa (Hook.) H. & A. ssp.
glandulosa

Layia platyglossa (F. & M.) Gray ssp.

NA

CR

campestris Keck

NA

MI

RO

CR

CA

CL

GU

Layia platyglossa (F. & M.) Gray ssp.
platyglossa

Lepidospartum squamatum (Gray)

NA

RO

Gray

NA

CR

Madia exigua (Sm.) Gray

NA

CR

CA

Madia gracilis (Sm.) Keck ssp. gracilis

NA

CR

CA

Madia sativa Mol.

IN

5

CR

CA

CL

Malacothrix clevelandii Gray
Malacothrix coulteri Harv. & Gray

NA

GU

var. cognata Jeps.

NA

RO

CR

Malacothrix “B” (see note 1)
Malacothrix foliosa Gray (Los Coro-

EN

20

nados)

EN

CL

Malacothrix incana (Nutt.) T. & G.

NA

MI

RO

CR

NI

76

Malacothrix indecora Greene

EN

MI

CR

Malacothrix “A” (see note 1)

EN

BA

Malacothrix “C” (see note 1 )
Malacothrix saxatilis (Nutt.) T. & G.

EN

NI

var. implicata (Eastw.) Hall
Malacothrix saxatilis (Nutt.) T. & G.

EN

MI

RO

CR

AN

NI

var. tenuifolia (Nutt.) Gray

NA

CA

Malacothrix similis Davis & Raven

NA

18

19

Malacothrix squalida Greene
Matricaria matricarioides (Less.) Por-

EN

CR

AN

ter

IN

RO

CR

CA

GU

Micropus californicus F. & M.
Microseris douglasii (DC.) Sch.-Bip.

NA

RO

CR

ssp. douglasii

Microseris douglasii (DC.) Sch.-Bip.

NA

81

NI

CL

ssp. platycarpha (Gray) Chamb.
Microseris douglasii (DC.) Sch.-Bip.

NA

CA

CL

ssp. tenella (Gray) Chamb.

NA

MI

RO

CR

8 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Microseris elegans Greene ex Gray

NA

Ml

66

CR

CL

Microseris heterocarpa (Nutt.) Chamb.

NA

RO

CR

CA

CL

GU

Microseris linearifolia (DC.) Sch.-Bip.

NA

RO

CR

AN

NI

72

CA

CL

GU

Pentachaeta lyonii Gray

NA

CA

Perezia microcephala (DC.) Gray

NA

RO

CR

CA

Perityle emoryi Torr. in Emory

NA

RO

CR

AN

BA

CA

CL

GU

Perityle incana Gray

EN

GU

Picris echioides L.

IN

CA

Pluchea odorata (L.) Cass.

NA

CR

CA

Pluchea sericea (Nutt.) Cov.
Psilocarphus tenellus Nutt. var. tenel-

NA

CA

lus

NA

RO

CR

CA

CL

Raphinesquia californica Nutt.

NA

RO

CR

AN

BA

CA

CL

Senecio aphanactis Greene

NA

RO

CR

CA

Senecio douglasii DC. ssp. douglasii

NA

CR

CA

CL

Senecio lyonii Gray

NA

CA

CL

Senecio palmeri Gray

EN

GU

Senecio vulgaris L.

IN

MI

CR

AN

NI

CA

CL

Silybum marianum (L.) Gaertn.

IN

RO

CR

BA

CA

Solidago californica Nutt.

NA

RO

CR

CA

Sonchus asper (L.) Hill

IN

MI

RO

CR

AN

NI

BA

CA

CL

Sonchus oleraceus L.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Sonchus tenerrimus L.

IN

31

31

NI

72

CA

CL

GU

Stephanomeria blairii M. & J.

EN

CL

Stephanomeria cichoriacea Gray

NA

RO

CR

Stephanomeria diegensis Gottlieb
Stephanomeria exigua Nutt. ssp. coro-

NA

RO

37

CA

CL

37

naria (Greene) Gottlieb

NA

MI

5

CR

Stephanomeria guadalupensis Bdg.
Stephanomeria virgata Benth. ssp. vir-

EN

GU

gata

NA

42

RO

CR

CA

CL

Sty locline gnaphalioides Nutt.

NA

RO

CR

CA

Taraxacum laevigatum (Willd.) DC.

IN

CR

CA

Taraxacum officinale Wiggers
Thelesperma megapotamicum

IN

CR

CA

(Spreng.) Kuntz

NA

CA

Tragopogon porrifolius L.

IN

CR

Venegasia carpesioides DC.

NA

RO

CR

Xanthium spinosum L.

Xanthium strumarium L. var. cana-

IN

MI

81

CR

BA

CA

dense (Mill.) T. & G.

NA

41

CA

Bataceae

Bat is maritima L.

NA

CL

Berberidaceae

Berberis pinnata Lag. ssp. insularis

Munz

EN

RO

CR

AN

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 9

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Boraginaceae

Amsinckia intermedia F. & M.

NA

MI

RO

CR

AN

BA

CA

CL

8

Amsinckia menziesii (Lehm.) Nels. &

Macbr.

NA

CR

CA

GU

Amsinckia spectabilis F. & M. var. ni-

colai (Jeps.) Jtn. ex Munz

EN

MI

NI

CL

Amsinckia spectabilis F. & M. var.
spectabilis

Cryptantha clevelandii Greene var.

NA

MI

RO

CR

AN

NI

BA

CL

96

clevelandii

NA

MI

RO

CR

AN

NI

BA

CA

CL

28

Cryptantha clevelandii Greene var. flo-

rosa Jtn.

NA

MI

RO

CR

AN

NI

BA

CA

Cryptantha foliosa (Greene) Greene

EN

GU

Cryptantha intermedia (Gray) Greene
Cryptantha leiocarpa (F. & M.)

NA

31

28

CA

CL

Greene

NA

42

RO

41

Cryptantha maritima (Greene) Greene

NA

NI

BA

CA

CL

GU

Cryptantha micromeres (Gray) Greene
Cryptantha microstachys (Greene ex

NA

RO

CR

CA

Gray) Greene

Cryptantha muricata (H. & A.) Nels.

NA

CA

& Macbr. var. jonesii (Gray) Jtn.

NA

CR

31

Cryptantha traskiae Jtn.

EN

NI

CL

Harpagonella palmeri Gray
Heliotropium curassavicum L. ssp.

NA

CA

GU

oculatum (Heller) Thome

NA

MI

RO

CR

AN

NI

CA

CL

Pectocarya linearis DC. ssp. ferocula
(Jtn.) Thome

Pectocarya penicellata (H. & A.)

NA

CR

29

CA

CL

GU

A. DC.

NA

31

41

AN

6

50

Pectocarya recurvata Jtn.
Plagiobothrys californicus (Gray)

NA

GU

Greene var. californicus
Plagiobothrys californicus (Gray)

NA

GU

Greene var. fulvescens Jtn.
Plagiobothrys californicus (Gray)

NA

MI

RO

CR

AN

66

Greene var. gracilis Jtn.

NA

31

CR

AN

CA

CL

Plagiobothrys canescens Benth.

NA

RO

CR

CA

CL

Brassicaceae

Arabis glabra (L.) Bemh.

IN

CR

Arabis hojfmannii (Munz) Roll.

EN

RO

CR

Athysanus pusillus (Hook.) Greene

NA

CR

CA

Brassica geniculata (Desf.) J. Ball

IN

MI

RO

CR

NI

CA

CL

Brassica kaber (DC.) L.C. Wheeler

IN

CR

CA

Brassica nigra (L.) Koch in Rohling
Brassica rapa L. ssp. sylvestris (L.)

IN

81

CR

NI

BA

CA

CL

GU

Janchen

IN

MI

81

CR

CA

CL

40

10 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San Santa Santa San Santa Santa San Isla

Occur- Miguel Rosa Cruz Anacapa Nicolas Barbara Catalina Clemente Guada-
Taxon rence Island Island Island Island Island Island Island Island lupe

Cakile edentula (Bigel.) Hook. var.
edentula

Cakile maritima Scop. ssp. maritima
Capsella bursa- pastoris (L.) Medic.
Cardamine californica (Nutt.) Greene
Cardamine oligosperma Nutt.

Cardaria draba (L.) Desv.

Caulanthus inflatus Wats.

Caulanthus lasiophyllus (H. & A.)
Pays.

Deseurainia pinnata (Walt.) Britt, ssp.

menziesii (DC.) Detl.

Dithyrea maritima A. Davids.

Draba cuneifolia Nutt, ex T. & G. var.

integrifolia Wats.

Erysimum ammophilum Heller
Erysimum cheiri (L.) Crantz
Erysimum insulare Greene
Erysimum moram Roll.

Hutchinsia procumbens (L.) Desv.
Lepidium lasiocarpum Nutt, ex T. &
G. var. lasiocarpum
Lepidium lasiocarpum Nutt, ex T. &
G. var. latifolium C.L. Hitchc.
Lepidium latipes Hook.

Lepidium nitidurn Nutt. var. nitidum
Lepidium oblongum Small
Lepidium virginicum L. var. pubescens
(Greene) Thell.

Lepidium virginicum L. var. robinsonii
(Thell.) C.L. Hitchc.

Lobularia maritima (L.) Desv.
Nasturtium officinale R. Br.

Raphanus raphanistrum L.

Raphanus sativus L.

Sibara filifolia (Greene) Greene
Sisymbrium altissimum L.

Sisymbrium irio L.

Sisymbrium officinale (L.) Scop.
Sisymbrium orientale L.

Thysanocarpus curvipes Hook. var.
curvipes

Thysanocarpus curvipes Hook. var.

elegans (F. & M.) Rob. in Gray
Thysanocarpus erectus Wats.
Thysanocarpus laciniatus Nutt, ex T.

& G. var. conchuliferus (Greene)
Jeps.

NA

MI

RO

81

IN

MI

RO

CR

AN

IN

MI

RO

CR

NA

MI

RO

CR

81

IN

CR

IN

CR

IN

CR

NA

MI

RO

CR

AN

NA

RO

CR

AN

NA

MI

NA

NA

RO

IN

NA

MI

RO

AN

EN

NA

MI

RO

CR

AN

NA

MI

RO

CR

26

NA

NA

CR

NA

44

RO

CR

AN

IN

MI

CR

AN

NA

41

NA

CR

IN

IN

MI

CR

IN

RO

IN

RO

CR

EN

CR

IN

IN

AN

IN

31

CR

IN

NA

CR

NA

NA

EN

CR

NI

NI

CA

CL

CA

CL

GU

CA

31

28

BA

CA

CL

GU

CA

CL

GU

NI

CA

CA

GU

NI

BA

GU

NI

CA

CL

GU

GU

CA

CL

NI

BA

CA

CL

GU

NI

CA

CL

GU

CA

CL

NI

NI

CA

CA

CA

CL

CA

CL

CA

CA

CA

CL

GU

CA

CA

CA

GU

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 11

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Thysanocarpus laciniatus Nutt, ex T.

& G. var. crenatus (Nutt.) Brewer
Thysanocarpus laciniatus Nutt, ex T.

NA

RO

CR

& G. var. laciniatus

NA

CR

CA

CL

Thysanocarpus laciniatus Nutt, ex T.

& G. var. ramosus (Greene) Munz

EN

RO

CR

Tropidocarpum gracile Hook.

NA

CA

CL

Cactaceae

Bergerocactus emoryi (Emgelm.) Britt.

& Rose

NA

CA

CL

Marnmillaria blossfeldiana Bodeker

var. shurliana Gates

NA

GU

Opuntia ficus-indica (L.) Mill.
Opuntia littoralis (Engelm.) Ckll. var.

IN

4

81

CA

littoralis

NA

MI

RO

CR

AN

NI

BA

CA

CL

Opuntia oricola Philbrick

NA

MI

72

4

AN

NI

BA

CA

CL

Opuntia prolifera Engelm.

NA

RO

CR

AN

NI

BA

CA

CL

GU

Callitrichaceae

Callitriche longipedunculata Morong
Callit riche marginata Torr. var. mar-

NA

CL

ginata

NA

CA

CL

Campanulaceae

Githopsis diffusa Gray ssp. diffusa (see

note 2)

NA

CR

GU

Lobelia erinus L.

IN

CR

Triodanis biflora (R. & P.) Greene

NA

RO

CR

CA

GU

Capparaceae

Cleome isomeris Greene

NA

RO

31

CA

CL

Caprifoliaceae

Lonicera hispidula (Lindl.) Dougl. ex
T. & G. var. vacillans Gray
Lonicera subspicata H. & A. var.

NA

31

5

CR

CA

CL

johnstonii Keck

NA

98

CR

CA

Sambucus mexicana Presl ex DC.
Syrnphoricarpos mollis Nutt, in

NA

RO

CR

CA

CL

T. & G.

NA

RO

CR

CA

Caryophyllaceae

Arenaria douglasii Fenzl. ex T. & G.

NA

RO

CR

CA

CL

Arenaria serpyllifolia L.

Cardionema ramosissima (Weinm.)

IN

CR

Nels. & Macbr.

NA

MI

RO

CR

Cerastium glomeratum Thuill.

IN

MI

RO

CR

CA

CL

GU

Herniaria cinerea DC.

IN

GU

Polycarpon depressum Nutt.

NA

CR

CA

96

Polycarpon tetraphyllum (L.) L.

IN

85

12 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Sagina decumbens (Ell.) T. & G. ssp.

occidentalis (Wats.) Crow

NA

MI

RO

CR

CA

Si/ene antirrhina L.

NA

42

RO

CR

CA

CL

GU

Silene gallica L.

Silene laciniata Cav. ssp. major

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Hitchc. & Maguire

NA

MI

RO

CR

AN

31

CL

Silene multinervia Wats.

NA

31

31

CR

31

CA

Spergula arvensis L.

Spergularia bocconii (Scheeie) Fou-

IN

RO

46

CA

caud

IN

CR

CA

CL

Spergularia macrotheca (Homem.)

Heynh. ssp. macrotheca

NA

MI

RO

CR

AN

NI

BA

CA

CL

GU

Spergularia marina (L.) Griseb.

NA

RO

CR

CA

CL

GU

Spergularia villosa (Pers.) Camb.

IN

RO

CA

CL

Stellaria media (L.) Vill.

IN

MI

RO

CR

CA

CL

Stellaria nit e ns Nutt.

NA

31

RO

CR

CA

GU

Chenopodiaceae

Aphanisma blitoides Nutt, ex Moq. in

DC.

NA

28

RO

CR

AN

NI

BA

CA

CL

GU

Atriplex argentea Nutt. ssp. expansa

(Wats.) Hall & Clem.

NA

RO

CR

CA

CL

Atriplex barclayana (Benth.) D. Dietr.

ssp. dilitata (Greene) Hail & Clem.
Atriplex barclayana (Benth.) D. Dietr.

NA

GU

ssp. palmeri (Wats.) Hall & Clem.

NA

GU

Atriplex californica Moq. in DC.

NA

MI

RO

CR

AN

NI

BA

CA

CL

GU

Atriplex coulteri (Moq.) D. Dietr.
Atriplex lentiformis (Torn) Wats. ssp.

NA

MI

RO

CR

AN

NI

CA

CL

breweri (Wats.) Hall & Clem.
Atriplex leucophylla (Moq. in DC.)

NA

31

CR

AN

31

CA

CL

D. Dietr.

NA

MI

RO

CR

AN

NI

CA

CL

Atriplex pacifica Nels.

Atriplex patula L. ssp. hastata (L.)

NA

CR

AN

31

CA

CL

Hall & Clem.

IN

MI

CR

NI

CA

Atriplex rosea L.

IN

CA

Atriplex semibaccata R. Br.

Atriplex serenana A. Nels. var. sere-

IN

MI

RO

CR

AN

NI

BA

CA

CL

nana

NA

46

46

CA

Atriplex watsonii A. Nels. in Abrams

NA

31

31

CR

NI

CA

CL

Bassia hyssopifolia (Pall.) Kuntze
Beta vulgaris L. ssp. maritima (L.) Ar-

NA

CA

CL

cang.

IN

MI

AN

CA

CL

Chenopodium album L.
Chenopodium ambrosioides L. var.

IN

5

CR

ambrosioides

IN

RO

CR

CA

Chenopodium berlandieri Moq. var.

sinuatum (J. Murr.) H.A. Wahl

NA

RO

CR

AN

CA

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 13

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Chenopodium californicum (Wats.)

Wats.

NA

MI

RO

CR

AN

NI

BA

CA

CL

Chenopodium multifidum L.

IN

RO

NI

Chenopodium murale L.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Monolepis nuttalliana (Schult.) Greene

NA

MI

RO

CR

CL

Salicornia subterminalis Parish

NA

MI

RO

CR

CA

CL

Salicornia virginica L.

NA

MI

RO

CR

AN

NI

CA

CL

Salsola iberica Sennen & Pau

IN

CR

NI

CA

CL

Suaeda californica Wats.

NA

MI

RO

CR

AN

NI

BA

CA

CL

GU

Cistaceae

Helianthemum greenei Rob.

EN

68

RO

CR

CA

Helianthemum scoparium Nutt.

NA

RO

CR

CA

Convolvulaceae

Calystegia macrostegia (Greene)

Brummitt ssp. amplissima Brum-

mitt

EN

NI

BA

CL

Calystegia macrostegia (Greene)

Brummitt ssp. cyclostegia (House)
Brummitt

NA

CA

Calystegia macrostegia (Greene)

Brummitt ssp. intermedia (Abrams)
Brummitt

NA

CA

Calystegia macrostegia (Greene)

Brummitt ssp. macrostegia (San
Martin Isl.)

EN

MI

RO

CR

AN

CA

GU

Calystegia soldanella (L.) R. Br.

NA

MI

RO

CR

58

CL

Convolvulus arvensis L.

IN

RO

CR

CA

Convolvulus simulans L.

NA

5

68

CL

Cressa truxillensis HBK. var. vallicola

(Heller) Munz

NA

MI

RO

CR

CA

CL

76

Cuscuta californica H. & A.

NA

RO

CL

Cuscuta ceanothi Behr.

NA

CR

Cuscuta corymbosa R. & P. var. gran-

diflora Engelm.

NA

GU

Cuscuta occidentalis Millsp.

NA

CA

Cuscuta salina Engelm. var. salina

NA

RO

CR

AN

Dichondra occidentalis House

NA

MI

RO

CR

CA

Ipomoea cairica (L.) Sweet

IN

CA

Ipomoea nil (L.) Roth

IN

CA

Comaceae

Cornus glabrata Benth.

NA

CA

Crassulaceae

Crassula aquatica (L.) Schoenl. in

Engl. & Prantl

NA

CA

Crassula erecta (H. & A.) Berger
Dudleya blochmanae (Eastw.) Moran

NA

MI

RO

CR

AN

NI

BA

CA

CL

GU

ssp. insularis (Moran) Moran

EN

RO

14 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Dudleya caespitosum (Haw.) Britt. &

Rose

NA

CR

AN

Dudleya candelabrum Rose

EN

RO

CR

Dudleya greenei Rose

EN

MI

RO

CR

AN

CA

Dudleya guadalupensis Moran

EN

GU

Dudleya hassei (Rose) Moran

EN

CA

Dudleya nesiotica (Moran) Moran

EN

CR

Dudleya traskiae (Rose) Moran

EN

BA

Dudleya virens (Rose) Moran

NA

NI

CA

CL

GU

Crossosomataceae

Crossosoma californicum Nutt.

NA

CA

CL

GU

Cucurbitaceae

Cucurbita foetidissima HBK.

NA

CR

Marah guadalupensis (Wats.) Greene

EN

GU

Marah macrocarpus (Greene) Greene

NA

Mi

RO

CR

AN

NI

BA

CA

CL

Elatinaceae

Elatine californica Gray

NA

CA

Ericaceae

Arbutus menziesii Pursh

NA

CR

Arctostaphylos catalinae P.V. Wells

EN

CA

Arctostaphylos confertiflora Eastw.
Arctostaphylos insularis Greene f. in-

EN

RO

sularis

EN

CR

Arctostaphylos insularis Greene f. pu-

bescens (Eastw.) P.V. Wells

EN

CR

Arctostaphylos tomentosa (Pursh)
Lindl. ssp. insulicola P.V. Wells
Arctostaphylos tomentosa (Pursh)

EN

RO

CR

Lindl. ssp. subcordata (Eastw.)
P.V. Wells

EN

RO

CR

Arctostaphylos viridissima (Eastw.)

McMinn

EN

CR

Comarostaphylis diversifolia (Parry)

Greene ssp. planifolia (Jeps.) Wal-
lace ex Thorne

NA

RO

CR

CA

Vaccinium ovatum Pursh

NA

RO

CR

Xylococcus bicolor Nutt.

NA

CA

Euphorbiaceae

Eremocarpus setigerus (Hook.) Benth.

NA

RO

CR

CA

CL

Euphorbia crenulata Engelm.

NA

CA

Euphorbia melanadenia Toit.

NA

95

Euphorbia misera Benth.

NA

CA

CL

GU

Euphorbia peplis L.

IN

RO

CR

CA

CL

Euphorbia pondii Millsp.

Euphorbia serpyllifolia Pers. var. ser-

NA

GU

pyllifolia

NA

CA

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 15

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Euphorbia spathulata Lam.

NA

CA

CL

Ricinus communis L.

IN

41

NI

CA

CL

Fabaceae

Acacia decurrens Willd.

IN

CA

Acacia malanoxylon R. Br.

IN

CR

CA

Astragalus curtipes Gray
Astragalus didymocarpus H. & A. ssp.

NA

MI

RO

didymocarpus

NA

MI

RO

CR

AN

CA

CL

Astragalus gambelianus Sheid.

NA

CR

CA

Astragalus miguelensis Greene

EN

MI

RO

CR

AN

CL

Astragalus nevinii Gray

EN

CL

Astragalus traskiae Eastw.
Astragalus trichopodus (Nutt.) Gray

EN

NI

BA

ssp. leucopsis (T. & G.) Thome
Astragalus trichopodus (Nutt.) Gray

NA

RO

CR

AN

CA

ssp. trichopodus

NA

CA

Coro nil la valentina L.

IN

CA

Cystisus linifolius (L.) Lam.

IN

CA

Cytisus monspessulanus L.

Lathyrus laetiflorus Greene ssp. alefel-

IN

CA

dii (White) Brads.

Lathyrus laetiflorus Greene ssp. bar-

NA

CA

barae (White) C.L. Hitchc.

NA

RO

CR

AN

CA

CL

Lathyrus tingitanus L.

Lathyrus vestitus Nutt, ex T. & G. ssp.

IN

CA

vestitus

NA

RO

CR

81

Lotus argophyllus (Gray) Greene ssp.

adsurgens (Dunkle) Raven
Lotus argophyllus (Gray) Greene ssp.

EN

CL

niveus (Greene) Munz
Lotus argophyllus (Gray) Greene ssp.

EN

CR

ornithopus (Greene) Raven

EN

NI

BA

CA

CL

GU

Lotus corniculatus L.

IN

CR

CA

Lotus grandiflorus (Benth.) Greene

var. grandiflorus

NA

RO

CR

CA

GU

Lotus hamatus Greene

NA

RO

CR

CA

CL

Lotus heermannii (Dur. & Hilg.)

Greene ssp. heermannii

NA

70

Lotus humistratus Greene

NA

CR

CA

Lotus micranthus Benth.

NA

CR

Lotus purshianus (Benth.) Clem. &
Clem. ssp. purshianus
Lotus salsuginosus Greene ssp. salsu-

NA

CR

CA

ginosus

NA

MI

RO

CR

AN

CA

Lotus scoparius (Nutt, in T. & G.) Ott-
ley var. dendroideus (Greene) Ottley
Lotus scoparius (Nutt, in T. & G.) Ott-

NA

MI

RO

CR

AN

CA

CL

ley ssp. scoparius

NA

MI

RO

CR

AN

CA

16 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Lotus scoparius (Nutt, in T. & G.) Ott-

ley ssp. traskiae (Eastw. ex Noddin
in Abrams) Raven

EN

CL

Lotus scoparius (Nutt, in T. & G.) Ott-
ley var. veatchii (Greene) Ottley
Lotus strigosus (Nutt, in T. & G.)

NA

MI

CR

Greene ssp. strigosus

NA

MI

RO

CR

AN

CA

CL

Lotus subpinnatus Lag.

NA

RO

CR

AN

CA

Lupinus agardhianus Heller

NA

RO

CR

CA

CL

Lupinus albifrons Benth. ssp. albifrons
Lupinus albifrons Benth. var. douglasii

NA

MI

RO

CR

AN

NI

CA

(J.G. Agardh) C.P. Sm.

NA

MI

RO

CR

NI

31

Lupinus arboreus Sims

Lupinus bicolor Lindl. ssp. microphyl-

NA

MI

RO

12

lus (Wats.) D. Dunn
Lupinus bicolor Lindl. ssp. piper-

NA

MI

RO

CR

AN

NI

CA

CL

GU

smithii (Heller) D. Dunn
Lupinus bicolor Lindl. ssp. tridentatus

NA

86

(Eastw. ex C.P. Sm.) D. Dunn

NA

CR

Lupinus bicolor Lindl. ssp. umbellatus
(Greene) D. Dunn
Lupinus concinnus J.G. Agardh ssp.

NA

MI

RO

CR

47

CA

CL

concinnus

NA

31

CR

CA

Lupinus densiflorus Benth. var. palus-

tris (Kell.) C.P. Sm.

NA

RO

CR

Lupinus guadalupensis Greene

EN

CL

GU

Lupinus hirsutissimus Benth.
Lupinus latifolius J.G. Agardh ssp.

NA

RO

CR

CA

CL

dudleyi (C.P. Sm.) Kenney & D.

Dunn

NA

91

Lupinus nanus Dougl. in Benth. ssp.

nanus

NA

41

Lupinus niveus Wats.

EN

GU

Lupinus polycarpus Greene

NA

RO

Lupinus succulentus Dougl. ex Koch

NA

MI

RO

CR

AN

CA

CL

Lupinus truncatus Nutt, ex H. & A.
Medicago polymorpha L. var. brevispi-

NA

RO

CR

AN

CA

CL

na (Benth.) Heyn.

Medicago polymorpha L. var. poly-

IN

RO

CR

CA

CL

GU

morpha

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Medicago sativa L.

IN

42

RO

CR

AN

NI

CA

CL

Melilotus alba Medicus

IN

CR

NI

CA

CL

Melilotus indica (L.) All.

Pickeringia montana Nutt. ssp. mon-

IN

MI

RO

CR

AN

NI

CA

CL

GU

tana

NA

CR

Spartium junceum L.

IN

NI

CA

Trifolium albopurpureum T. & G.
Trifolium amplectens T. & G. var.

NA

RO

CR

NI

CA

amplectens

NA

MI

RO

CR

81

28

CA

CL

GU

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 17

Taxon

Occur-

rence

San

Miguel

Island

Santa

Rosa

Island

Santa

Cruz

Island

Anacapa

Island

San

Nicolas

Island

Santa

Barbara

Island

Santa San

Catalina Clemente
Island Island

Isla

Guada-

lupe

Trifolium amplectens T. & G. var.

truncatum (Greene) Jeps.

NA

47

RO

CR

AN

CA

33

Trifolium barbigerum Torr.

NA

MI

RO

Trifolium ciliolatum Benth.

NA

RO

CR

CA

Trifolium depauperatum Desv.

NA

CR

NI

Trifolium fucatum Lindl. var. gambelii

(Nutt.) Jeps.

NA

MI

RO

CR

CA

CL

Trifolium gracilentum T. & G.

NA

MI

RO

CR

AN

NI

CA

CL

GU

Trifolium macraei H. & A.

NA

RO

CR

CA

Trifolium microcephalum Pursh

NA

MI

RO

CR

CA

CL

GU

Trifolium microdon H. & A. var. pi-

losum Eastw.

EN

41

NI

CA

Trifolium palmeri Wats.

EN

NI

BA

CA

CL

GU

Trifolium repens L.

IN

CA

Trifolium tridentatum Lindl. var. aci-

culare (Nutt.) McDer.

NA

MI

RO

CR

AN

NI

BA

CA

CL

Trifolium tridentatum Lindl. var. tri-

dentatum

NA

MI

RO

CR

AN

NI

BA

CA

CL

Trifolium variegatum Nutt, in T. & G.

NA

CR

Vicia americana Muhl. ex Willd.

NA

MI

RO

CR

Vicia dasycarpa Ten.

IN

NI

CA

Vicia exigua Nutt, in T. & G.

NA

42

RO

CR

AN

CA

CL

GU

Vicia hassei Wats.

NA

RO

CR

AN

NI

57

CL

GU

Vicia villosa Roth

IN

NI

Fagaceae

Quercus agrifolia Nee var. agrifolia

NA

RO

CR

Quercus chrysolepis Liebm.

NA

CR

CA

CL

Quercus douglasii H. & A.

NA

CR

CA

Quercus dumosa Nutt.

NA

RO

CR

CA

Quercus engelmannii Greene

NA

CA

Quercus lobata Nee

NA

CR

CA

Quercus x macdonaldii Greene

EN

RO

CR

CA

Quercus x morehus Kell.

EN

CR

Quercus tomentella Englem.

EN

RO

CR

AN

CA

CL

GU

Quercus mslizenii A. DC. var. frutes-

cens Englem.

NA

CR

Frankeniaceae

Frankenia grandifolia Cham. &

Schlecht. var. grandifolia

NA

MI

RO

CR

AN

NI

CA

CL

GU

Garryaceae

Garrya elliptica Dougl.

NA

CR

Gentianaceae

Centaurium davyi (Jeps.) Abrams

NA

RO

CR

CL

Centaurium exaltatum (Griseb.) W.

Wight

NA

5

CR

Centaurium muhlenbergii (Griseb.) W.

Wight

NA

52

18 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Centaurium venustum (Gray) Rob.

NA

CA

Geraniaceae

Erodium botrys (Cav.) Berlol.

IN

RO

CR

CL

Erodium cicutarium (L.) L’Her.
Erodium macrophyllum H. & A. var.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

californicum (Greene) Jeps.

NA

CR

Erodium moschatum (L.) L’Her.
Erodium obtusiplicatum (Maire, Weill-

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

er & Wilcz.) J.T. Howell

IN

RO

CA

Erodium texanum Gray

NA

CA

Geranium carolinianum L.

IN

RO

CR

CA

Geranium dissectum L.

IN

RO

Pelargonium x hortorum Bailey
Pelargonium peltatum (L.) L’Her. ex

IN

RO

CR

NI

CA

Ait.

IN

NI

Hydrophyllaceae

Emmenanthe pendulijlora Benth.
Eriodictyon traskiae Eastw. ssp. Iras-

NA

CR

CA

CL

GU

kiae

EN

CA

Eucrypta chrysanthemifolia (Benth.)

Greene var. chrysanthemifolia
Nemophila menziesii H. & A. ssp.

NA

MI

RO

CR

AN

CA

CL

GU

menziesii

NA

CA

Nemophila pedunculata Dougl. ex

Benth.

NA

MI

RO

CR

49

Phacelia cicutaria Greene ssp. hispida

(Gray) J. Beauchamp ex Thome

NA

RO

CR

AN

CA

CL

Phacelia cinerea Eastw. ex Macbr.

EN

NI

Phacelia distans Benth.

Phacelia divaricata (Benth.) Gray var.

NA

MI

RO

CR

AN

BA

CA

CL

insularis (Munz) Munz

EN

MI

RO

Phacelia floribunda Greene

EN

CL

GU

Phacelia grandiflora (Benth.) Gray

NA

RO

CR

CA

Phacelia lyonii Gray

EN

CA

CL

Phacelia phyllomanica Gray
Phacelia ramosissima Dougl. ex

EN

GU

Lehm. var. austrolitoralis Munz

NA

RO

CR

Phacelia ramosissima Dougl. ex

Lehm. var. montereyensis Munz

NA

MI

RO

Phacelia viscida (Benth.) Torr.
Pholistoma auritum (Lindl.) Lilja ex

NA

42

RO

CR

AN

CA

Lind.

NA

CR

BA

CA

CL

Pholistoma racemosum (Nutt.) Const.

NA

CR

BA

CA

CL

GU

Juglandaceae

Juglans californica Wats.

NA

CA

Lamiaceae

Lamium amplexicaule L.

IN

CR

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 19

San

Santa

Santa

San

Santa

Santa

San

Isia

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Lepechinia calycina (Benth.) Epl. in

Munz

NA

RO

BA

Lepechinia fragrans (Greene) Epl.

NA

RO

CR

CA

Marrubium vulgare L.

IN

MI

RO

CR

NI

CA

CL

Mentha citrata Ehrh.

IN

CA

Mentha spicata L.

IN

81

CA

Nepeta cataria L.

IN

CA

Pogogyne tenuiflora Gray

EN

GU

Salvia apiana Jeps. var. apiana

NA

CA

Salvia brandegei Munz

Salvia columbariae Benth. ssp. colum-

NA

RO

bariae

NA

RO

CR

CA

CL

Salvia mellifera Greene

NA

RO

CR

AN

CA

Satureja douglasii (Benth.) Briq.

NA

CA

Satureja palmeri (Gray) Briq.
Scutellaria tuberosa Benth. ssp. a us-

EN

GU

tralis Epl.

NA

CR

Stachys ajugoides Benth.

NA

52

Stachys bullata Benth.

NA

RO

CR

AN

Trichostema lanceolatum Benth.

NA

CA

Linaceae

Hesperolinon micranthum (Gray)

Small

NA

CA

Loasaceae

Mentzelia affinis Greene
Mentzelia micrantha (H. & A.)

NA

CR

CA

CL

T. & G.

NA

5

CR

CA

CL

GU

Lythraceae

Ammannia coccinea Rottb.

NA

CA

Lythrum californicum T. & G.

NA

CR

Malvaceae

Althaea rosea (L.) Cav.

IN

CA

Eremalche exilis (Gray) Greene
Lavatera assurgenti flora Kell, (see note

NA

5

CA

CL

3)

EN

MI

RO

31

NI

CA

CL

Lavatera lindsayi Moran

EN

GU

Lavatera occidentalis Wats.
Malacothamnus clementinus (M. & J.)

NA

GU

Keam.

EN

CL

Malacothamnus fasciculatus (Nutt.)

Greene ssp. catalinensis (Eastw.)
Thome

NA

CA

Malacothamnus fasciculatus (Nutt.)

Greene var. nesioticus (Rob.) Keam.

EN

CR

Malva parviflora L.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Malvella leprosa (Ortega) Krapovickas

NA

81

CA

CL

11

20 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Mliguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Sidalcea malvaeflora (DC.) Gray

ex Benth ssp. malvaeflora

NA

MI

RO

CR

Sphaeralcea palmeri Rose

EN

GU

Sphaeralcea sulphurea Wats.

EN

GU

Moraceae

Ficus carica L.

IN

MI

CR

CA

Myrtaceae

Eucalyptus globulus Labill.

IN

RO

CR

AN

NI

CA

Nyctaginaceae

Abronia latifolia Esch.

NA

MI

Abronia maritima Nutt, ex Wats.

NA

MI

RO

CR

AN

NI

CA

CL

Abronia umbellata Lam.

NA

MI

RO

CR

NI

CA

CL

Mirabilis californica Gray var. ca/ifor-

nica

NA

46

CR

AN

BA

CA

CL

Mirabilis heimerlii (Standi.) Macbr.

EN

GU

Oleaceae

Hesperelaea palmeri Gray

EN

GU

Olea europaea L.

IN

CR

Onagraceae

Camissonia californica (Nutt, ex
T. & G.) Raven

Camissonia cheiranthifolia (Homem.

NA

RO

CR

CA

ex Spreng.) Raim. in Engl. & Prantl
ssp. cheiranthifolia

NA

MI

RO

CR

NI

BA

CL

Camissonia cheiranthifolia (Homem.

ex Spreng.) Raim. in Engl. & Prantl
ssp. suffruticosa (Wats.) Raven

NA

NI

Camissonia guadalupensis (Wats.) Ra-
ven ssp. Clementina (Raven) Raven
Camissonia guadalupensis (Wats.) Ra-

EN

CL

ven ssp. guadalupensis

EN

GU

Camissonia hirtella (Greene) Raven

NA

CR

Camissonia ignota (Jeps.) Raven

NA

CA

Camissonia intermedia Raven
Camissonia micrantha (Homem. ex

NA

CR

CA

Spreng.) Raven

NA

MI

RO

CR

CA

CL

Camissonia robusta Raven

NA

MI

CR

CA

CL

GU

Camissonia strigulosa (F. & M.) Ra-

ven

NA

47

RO

Clarkia davyi (Jeps.) Lewis & Lewis
Clarkia epilobioides (Nutt.) Nels. &

NA

RO

Macbr.

NA

RO

CR

81

CA

CL

Clarkia prostrata Lewis & Lewis
Clarkia purpurea (Curt.) Nels. &

NA

RO

Macbr. ssp. quadrivulnera (Dougl. in
Lindl.) Lewis & Lewis

NA

RO

CR

CA

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 21

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Clarkia unguiculata Lindl.

Epilobium canum (Greene) Raven ssp.

NA

5

CR

CA

canum

NA

42

RO

CR

AN

CA

CL

Epilobium ciliatum Raf. ssp. ciliatum

NA

MI

CR

CA

Epilobium foliosum (T. & G.) Suksd.

NA

GU

Gaura sinuata Nutt, ex Ser. in DC.

IN

CA

Ludwigia peploides (HBK.) Raven ssp.
peploides

Oenothera elata HBK. ssp. hirsutissi-

NA

CR

ma (Gray ex Wats.) Dietrich

NA

CR

Orobanchaceae

Orobanche bulbosa G. Beck
Orobanche californica Cham. &

NA

RO

CR

CA

Schlecht. ssp. grandis Heckard

NA

RO

Orobanche fasciculata Nutt.
Orobanche parishii (Jeps.) Heckard

NA

81

RO

CR

CA

ssp. brachyloba Heckard
Orobanche uniflora L. ssp. occidentals

NA

MI

RO

CR

NI

CA

(Greene) Abrams ex Ferris

NA

81

CR

Oxalidaceae

Oxalis albicans HBK. ssp. californica
(Abrams) Eiten

Oxalis albicans HBK. ssp. pilosa

NA

CR

CA

(Nutt.) Eiten

NA

CR

Oxalis corniculata L.

IN

CR

CA

Oxalis pes-caprae L.

IN

CA

CL

Papaveraceae

Dendromecon rigidus Benth. ssp. har-
fordii (Kell.) Raven
Dendromecon rigidus Benth. ssp.

EN

RO

CR

rhamnoides (Greene) Thome

EN

CA

CL

Eschscholzia californica Cham. var.
californica

Eschscholzia californica Cham. var.

NA

MI

RO

CR

CA

GU

maritima (Greene) Jeps.
Eschscholzia californica Cham. var.

NA

MI

RO

CR

peninsularis (Greene) Munz

NA

CR

CA

Eschscholzia elegans Greene
Eschscholzia frutescens (Greene)

EN

GU

J.T. Howell

EN

GU

Eschscholzia palmeri Rose

EN

GU

Eschscholzia ramosa (Greene) Greene

NA

RO

CR

26

73

BA

CA

CL

GU

Meconella denticulata Greene

NA

CR

Papaver californicum Gray

NA

CR

Papaver somniferum L.

IN

CA

Platystemon californicus Benth.

NA

MI

RO

CR

AN

NI

BA

CA

GU

Romneya coulteri Harv.

NA

CA

22 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Stylomecon heterophylla (Benth.)

G. Taylor

NA

MI

RO

CR

AN

BA

CA

CL

Pittosporaceae

Sollya heterophylla Lindl.

IN

CA

Plantaginaceae

Planlago bigelovii Gray ssp. californica

(Greene) Bassett

NA

MI

RO

Plant ago coronopus L.

IN

CA

Plantago erecta Morris ssp. erecta
Plantago hirtella HBK. ssp. galleot-

NA

42

RO

CR

AN

CA

CL

liana (Dene.) Thome

NA

42

RO

CR

Plantago lanceolata L.

IN

CR

CL

Plantago major L.

Plantago maritima L. var. californica

IN

CR

CA

(Fern.) Pilg.

NA

RO

Planlago ovata Forssk.

IN

RO

AN

NI

BA

CA

CL

GU

Platanaceae

Platanus racemosa Nutt.

NA

CR

CA

Plumbaginaceae

Armeria maritima (Mill.) Willd. ssp.

californica (Boiss.) G.FI.M. Lawr.
Limonium perezii (Stapf.) F.T. Hubb.

NA

RO

ex Bailey

IN

CA

CL

Limonium sinuatum (L.) Mill.

IN

CA

Polemoniaceae

Allophyllum gilioides (Benth.)

A. Grant & V. Grant

NA

GU

Allophyllum glutinosum (Benth.)

A. Grant & V. Grant

NA

CA

Eriastrum filifolium (Nutt.) Woot. &

Standi.

NA

CR

CA

CL

Gilia angelensis V. Grant

Gilia capitata Sims ssp. abrotanifolia

NA

CR

AN

CA

CL

(Nutt, ex Greene) V. Grant

NA

CR

CA

Gilia clivorum (Jeps.) V. Grant

NA

Ml

RO

CR

AN

CA

Gilia nevinii Gray

Gilia tenuiflora Benth. ssp. hoffmannii

EN

RO

CR

AN

NI

BA

CA

CL

GU

(Eastw.) A. Grant & V. Grant

EN

RO

Linanthus androsaceus (Benth.)

Greene ssp. luteus (Benth.) Mason
Linanthus bicolor (Nutt.) Greene ssp.

NA

81

5

CR

bicolor

NA

42

RO

31

CA

CL

Linanthus dianthiflorus (Benth.)

Greene ssp. dianthiflorus
Linanthus pygmaeus (Brand)

NA

CR

CA

J.T. Howell ssp. pygmaeus

EN

CL

GU

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 23

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Navarretia atractyloides (Benth.)

H. & A.

NA

RO

CR

CA

CL

Navarretia hamata Greene var. foli-

acea (Greene) Thome

NA

CA

Navarretia hamata Greene var.

hamata

NA

CR

CA

CL

96

Polygalaceae

Polygala californica Nutt.

NA

CR

Polygonaceae

Chorizanthe coriacea Goodm.
Chorizanthe staticoides Benth. ssp.

NA

RO

CR

CA

staticoides

NA

5

CR

CA

Chorizanthe wheeled Wats.

NA

RO

CR

Eriogonum arborescens Greene

EN

RO

CR

AN

Eriogonum cinereum Benth.
Eriogonum fasciculatum Benth. ssp.

NA

RO

fasciculatum

NA

CA

Eriogonum giganteum Wats. ssp. com-
pactum (Dunkle) Munz
Eriogonum giganteum Wats. ssp. for-

EN

BA

mosum (K. Bdg.) Raven

EN

CL

Eriogonum giganteum Wats. ssp. gi-
ganteum (see note 4)

Eriogonum grande Greene var. dunk-

EN

CA

lei Reveal

EN

MI

Eriogonum grande Greene ssp. grande
Eriogonum grande Greene ssp. rubes-

NA

RO

CR

AN

NI

CA

CL

cens (Greene) Munz
Eriogonum grande Greene ssp. timo-

EN

MI

RO

CR

AN

rum (Reveal) Munz

EN

NI

Eriogonum zapatoense Moran

EN

GU

Polygonum arenastrum Bor.
Polygonum argyrocoleon Steud. ex

IN

RO

CR

CA

CL

Kunze

IN

CR

CA

CL

Polygonum aviculare L.

IN

RO

CR

CA

CL

Pterostegia drymarioides F. & M.

NA

MI

RO

CR

AN

BA

CA

CL

GU

Rumex angiocarpus Murbeck

IN

CR

Rumex conglomeratus Murr.

IN

RO

CR

CA

Rumex crispus L.

IN

MI

RO

CR

AN

NI

CA

CL

Rumex fueginus Phil.

NA

MI

41

31

Rumex pulcher L.

IN

CR

Rumex salicifolius Weinm.

NA

MI

RO

CR

NI

CA

CL

Portulacaceae

Calandrinia brewed Wats.

Calandrinia ciliata (R. & P.) DC. var.

NA

RO

CR

menziesii (Hook.) Macbr.

NA

MI

RO

CR

AN

BA

CA

CL

GU

Calandrinia maritima Nutt.

NA

RO

CR

AN

BA

CA

CL

GU

24 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Calyptridium monandrum Nutt, in

T. & G.

NA

52

Claytonia perfoliata Donn var. parvi-
Jlora (Dougl. ex Hook.) Torn
Claytonia perfoliata Donn var. perfoli-

NA

CR

ata

NA

MI

RO

CR

AN

NI

BA

CA

CL

GU

Montia fontana L. ssp. amporitana

Sennen

NA

CR

31

Portulaca oleracea L.

IN

CR

CA

Talinum gnadalupense Dudl. in

D.S. Jordan

EN

GU

Primulaceae

Anagalis arvensis L.

IN

31

RO

CR

AN

CA

93

Anagalis minima (L.) Krause
Dodecatheon clevelandii Greene ssp.

NA

RO

insularis H.J. Thomps.

NA

RO

CR

AN

CA

CL

GU

Samolus parviflorus Raf.

NA

CR

Ranunculaceae

Clematis lasiantha Nutt, in T. & G.

NA

31

CR

Clematis ligusticifolia Nutt, in T. & G.

NA

RO

CR

CA

Delphinium kinkiense Munz

EN

CL

Delphinium parryi Gray ssp. parryi
Delphinium variegatum T. & G. ssp.

NA

MI

RO

CR

AN

CA

CL

thornei Munz

EN

CL

Myosurus minimus L. var. filiformis

Greene

NA

GU

Ranunculus californicus Benth. ssp.
californicus

Ranunculus californicus Benth. var.

NA

MI

RO

CR

cuneatus Greene

NA

MI

31

CR

31

Ranunculus hebecarpus H. & H.

NA

CA

GU

Resedaceae

Oligomeris linifolia (Vah.) Macbr.

NA

MI

RO

CR

AN

NI

BA

CA

CL

GU

Reseda odorata L.

IN

31

CA

Rhamnaceae

Ceanothus arboreus Greene

EN

RO

CR

CA

Ceanothus crassifolius Torr.
Ceanothus megacarpus Nutt. ssp. in-

NA

GU

sularis (Eastw.) Raven
Ceanothus megacarpus Nutt. ssp.

EN

42

RO

CR

AN

CA

CL

megacarpus

NA

CR

CA

CL

Rhamnus californica Esch. ssp. califor-

nica

NA

CR

Rhamnus pirifolia Greene

EN

42

RO

CR

CA

CL

GU

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 25

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Rosaceae

Adenostoma fasciculatum H. & A. var.

fasciculatum

NA

RO

CR

CA

CL

Alchemil/a occidentalis Nutt.

NA

RO

CR

CA

CL

GU

Cercocarpus betuloides Nutt, ex T. &
G. ssp. betuloides

Cercocarpus betuloides Nutt, ex T. &

NA

31

CR

CA

G. ssp. blancheae (C.K. Schneid.)
Thome

NA

RO

CR

CA

Cercocarpus traskiae Eastw.
Heteromeles arbutifolia (Ait.) M.

EN

CA

Roem.

NA

MI

RO

CR

AN

31

CA

CL

GU

Holodiscus discolor (Pursh) Maxim,
ssp. discolor

Lyonothamnus Jloribundus Gray ssp.

NA

CR

CA

asplenifolius (Greene) Raven (see
note 5)

EN

RO

CR

CL

Lyonothamnus Jloribundus Gray ssp.
Jloribundus

Potentilla egedii Wormsk. var. grandis

EN

CA

(Rydb.) J.T. Howell
Potentilla glandulosa Lindl. ssp. glan-

NA

MI

CR

dulosa

NA

CA

Prunus lyonii (Eastw.) Sarg.

NA

RO

CR

AN

CA

CL

Prunus persica (L.) Batsch

IN

CA

Rosa californica Cham. & Schlecht.

NA

RO

CR

CA

Rubus procerus P.J. Muell.

IN

CA

Rubus ursinus Cham. & Schlecht.

NA

MI

RO

CR

26

CA

Rubiaceae

Galium angulosum Gray
Galium angustifolium Nutt, ex T. &

EN

GU

G. ssp. angustifolium
Galium angustifolium Nutt, ex T. &

NA

CA

G. ssp. foliosum (Hilend & Howell)
Dempst. & Steb.

EN

RO

CR

AN

Galium aparine L.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Galium buxifolium Greene
Galium californicum H. & A. ssp.

EN

MI

RO

CR

flaccidum (Greene) Dempst.
Galium californicum H. & A. ssp.

NA

CR

miguelense (Greene) Dempst. &
Steb.

EN

MI

RO

Galium catalinense Gray ssp. acris-
pum Dempst.

Galium catalinense Gray ssp. catali-

EN

CL

nense

EN

CA

Galium nuttallii Gray ssp. insulare

Ferris

EN

RO

CR

31

CA

26 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Galium porrigens Dempst. var. porn-

gens

NA

RO

CR

22

Rutaceae

Rut a chalepensis L.

IN

CR

CA

GU

Salicaceae

Populus fremontii Wats. ssp. fremontii

NA

31

CR

CA

Populus trichocarpa T. & G.

NA

RO

CR

CA

Populus x parryi Sarg.

NA

NI

CA

Sal ix hindsiana Benth. var. hindsiana

NA

CR

CA

Salix laevigata Bebb. var. laevigata

NA

CR

CA

Sa/ix lasiandra Benth. var. lasiandra

NA

CR

Salix lasiolepis Benth. var. lasiolepis

NA

Ml

RO

CR

NI

CA

Saururaceae

Anemopsis californica (Nutt.) H. & A.

NA

31

31

CR

NI

CA

76

Saxifragaceae

Heuchera maxima Greene

EN

RO

CR

AN

Jepsonia malvaefolia (Greene) Small
Lithophragma affine Gray ssp. mix-

EN

RO

CR

NI

CA

CL

GU

turn R.L. Taylor

NA

CA

Lithophragma cymbalaria T. & G.

NA

RO

CR

Lithophragma maximum Bacig.
Rihes malvaceum Sm. in Rees var.

EN

CL

malvaceum

NA

RO

CR

AN

CL

Ribes menziesii Pursh var. menziesii
Ribes menziesii Pursh var. thacheria-

NA

CR

num Jeps.

EN

CR

Ribes sanguineum Pursh

NA

GU

Ribes viburnifolium Gray

NA

CA

Saxifraga californica Greene

NA

RO

CR

Scrophulariaceae

Antirrhinum kelloggii Greene
Antirrhinum kingii Wats. var. watsoni

NA

CR

CA

(Vasey & Rose) Munz

NA

GU

Antirrhinum multiflorum Penn.
Antirrhinum nuttallianum Benth. in

NA

CR

DC.

NA

MI

RO

CR

AN

CA

CL

GU

Castilleja affinis H. & A. ssp. affinis

NA

42

RO

CR

AN

CA

Castilleja foliolosa H. & A.

NA

CA

Castilleja fruticosa Moran

EN

GU

Castilleja grisea Dunkle

EN

CL

Castilleja guadalupensis Bdg.

EN

GU

Castilleja hololeuca Greene

EN

MI

RO

CR

AN

Castilleja mollis Penn.

Collinsia heterophylla Buist ex Grah.

NA

MI

RO

ssp. heterophylla

NA

31

RO

31

CL

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 27

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Diplacus longiflorus Nutt. ssp. longi-

florus

NA

RO

CR

Diplacus parviflorus Greene

EN

RO

CR

AN

CL

Diplacus puniceus Nutt.

NA

CA

Galvezia speciosa (Nutt.) Gray

NA

BA

CA

CL

GU

Keckiella cordifolia (Benth.) Straw

NA

RO

CR

AN

CA

CL

Linaria bipartita Willd.

Linaria canadensis (L.) Dum.-Cours.

IN

CA

var. texana (Scheele) Penn.

NA

MI

RO

CR

AN

CA

CL

GU

Mimulus brandegei Penn.

EN

CR

Mimulus brevipes Benth.

NA

67

CA

Mimulus cardinalis Dougl. ex Benth.
Mimulus jloribundus Dougl. ex Lindl.

NA

CR

CA

var. jloribundus

Mimulus guttatus Fisch. ex DC. ssp.

NA

RO

41

CA

CL

guttatus

Mimulus guttatus Fisch. ex DC. ssp.

NA

RO

CR

CA

CL

lit! oralis Penn.

NA

RO

Mimulus guttatus Fisch. ex DC. ssp.

micranthus (Heller) Munz

NA

CR

Mimulus latifolius Gray

EN

GU

Mimulus nasutus Greene
Mimulus traskiae Grant in Millsp. &

NA

CR

Nutt.

EN

CA

Orthocarpus attenuatus Gray
Orthocarpus densijlorus Benth. var.

NA

81

GU

densijlorus

Orthocarpus purpurascens Benth. var.

NA

MI

RO

CR

NI

pallidus Keck

Orthocarpus purpurascens Benth. var.

NA

MI

RO

CR

CA

purpurascens

NA

MI

RO

CR

CA

8

Scrophularia californica Cham. &
Schlecht. ssp. californica
Scrophularia villosa Penn, in Millsp. &

NA

RO

Nutt.

EN

CA

CL

GU

V erbascum thapsus L.

IN

CR

Solanaceae

Datura wrightii Regel

NA

RO

CR

CA

Lycium brevipes Benth. var. brevipes
Lycium brevipes Benth. var. hassei

NA

CL

(Greene) C.L. Hitchc.

EN

CA

CL

Lycium californicum Nutt.

NA

AN

NI

BA

CA

CL

GU

Lycium fremontii Gray

NA

RO

GU

Lycium verrucosum Eastw.

Nicotiana attenuata Torr. ex Wats, in

EN

NI

King

Nicotiana bigelovii (Torr.) Wats. var.

NA

GU

bigelovii

NA

CA

28 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Nicotiana clevelandii Gray

NA

CR

CA

Nicotiana glauca Grab.

IN

CA

GU

Petunia parvi flora Juss.

NA

RO

Solanum douglasii Dunal in DC.

NA

MI

RO

CR

CA

CL

GU

Solanum elaeagnifolium Cav.

IN

CR

CA

Solanum nodiflorum Jacq.

IN

NI

CL

GU

Solatium sarrachoides Sendt. ex Mart.

IN

CR

Solanum wallacei (Gray) Parish ssp.

clokeyi (Munz) Thorne
Solanum wallacei (Gray) Parish ssp.

EN

RO

CR

wallacei

EN

CA

GU

Tamaricaceae

Tamarix tetrandra Pallas

IN

CR

AN

Tropaeolaceae

Tropaeolum majus L.

IN

CA

Urticaceae

Hesperocnide tenella Torr.

NA

CR

BA

CA

CL

GU

Parietaria hespera Hinton

NA

MI

RO

CR

AN

NI

BA

CA

CL

GU

Soleirolia soleirolii (Req.) Dandy
Urtica dioica L. ssp. holoserica (Nutt.)

IN

NI

Thorne

NA

CR

AN

CA

Urtica urens L.

IN

RO

CR

CA

Valerianaceae

Centranthus ruber (L.) DC.

IN

CR

CA

Verbenaceae

Lippia nodiflora (L.) Michx. var. rosea

(D. Don) Munz

IN

CR

CA

Verbena bracteata Lag. & Rodr.

NA

CA

Verbena lasiostachys Link

NA

42

81

CR

CA

CL

Verbena robusta Greene

NA

31

RO

CR

CA

Violaceae

Viola pedunculata T. & G. ssp. pedun-

culata

NA

RO

CR

CA

CL

Viscaceae

Phoradendron bolleanum (Seem.)

Eichler ssp. densum (Torr.) Wiens

NA

GU

Vitaceae

Vitis girdiana Munson

NA

CA

Monocotyledons

Arecaceae

Erythaea edulis (Wendl.) Wats.

EN

GU

Cyperaceae

Carex barbarae Dewey

NA

CR

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 29

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Carex globosa Boott.

NA

RO

CR

31

Carex gracilior Mkze.

NA

RO

CR

Carex montereyensis Mkze.

NA

CR

Carex pansa Bailey

NA

RO

Carex praegracilis W. Boott.

NA

RO

CR

CA

Carex rossii Boott.

NA

81

Carex sent a Boott.

NA

CR

Carex subbracteata Mkze.

NA

RO

CR

Carex triquetra Boott.

NA

31

CR

CA

Carex tumulicola Mkze.

NA

81

CR

CL

Cyperus alternifolius L.
Eleocharis macrostachya Britt, in

IN

CR

Small

NA

RO

29

NI

CA

CL

Scirpus americanus Pers. var. mono-

phyllus (Presl) Koyama

NA

RO

Scirpus californicus (C.A. Mey.) Steud.
Scirpus cernuus Vahl. ssp. californicus

NA

81

CR

(Torr.) Thome

NA

MI

RO

Scirpus microcarpus Presl

NA

CA

Scirpus olneyi Gray

NA

RO

NI

Scirpus robustus Pursh

NA

63

Iridaceae

Chasmanthe aethiopica (L.) N.E. Br.

IN

MI

Iris ochroleuca L.

IN

NI

Sisyrinchium bellum Wats.

NA

MI

RO

CR

CA

Juncaceae

Juncus acutus L. var. sphaerocarpus

Englem.

NA

CA

Juncus balticus Willd.

NA

MI

RO

CR

CA

Juncus bufonius L.

NA

MI

RO

CR

NI

CA

CL

GU

Juncus effusus L. var. bruneus Engelm.
Juncus effusus L. var. pacificus Fem.

NA

CR

& Wieg.

NA

81

CR

Juncus mexicanus Willd.

NA

MI

RO

CR

CA

Juncus patens E. Mey.

Juncus phaeocephalus Engelm. var.

NA

RO

CR

CL

phaeocephalus

NA

RO

Juncus textilis Buch.

NA

CA

Juncus xiphioides E. Mey.

NA

RO

CR

CA

Luzula subsessilis (Wats.) Buch.

NA

31

RO

CR

Liliaceae

Allium lacunosum Wats. var. lacuno-

sum

NA

RO

CR

Allium praecox Bdg.

NA

MI

RO

CR

CA

CL

Asparagus officinalis L.

Bloomeria crocea (Torr.) Cov. ssp.

IN

CA

crocea

NA

RO

CR

CA

30 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San

Santa

Santa

San

Santa

Santa

San

Isla

Occur-

Miguel

Rosa

Cruz

Anacapa

Nicolas

Barbara

Catalina Clemente

Guada-

Taxon

rence

Island

Island

Island

Island

Island

Island

Island

Island

lupe

Brodiaea jolonensis Eastw.

NA

81

RO

CR

CA

Brodiaea kinkiensis Niehaus
Calochortus albus Dougl. ex Benth.

EN

CL

var. albus

NA

RO

CR

CA

Calochortus catalinae Wats.

NA

RO

CR

CA

Calochortus luteus Dougl. ex Lindl.
Calochortus splendens Dougl. ex

NA

CR

31

Benth.

NA

CA

Chlorogalum pomeridianum (DC.)

Kunth

NA

RO

CA

Dichelostemma pulchellum (Salisb.)

Heller

NA

Ml

RO

CR

AN

NI

BA

CA

CL

GU

Lilium humboldtii Roezl & Leichtl.

ssp. ocellatum (Kell.) Thome

NA

RO

CR

Triteleia Clementina Hoov.

EN

CL

Triteleia guadalupensis Lenz
Zigadenus fremontii Torr. var. fre-

EN

GU

montii

NA

RO

CR

AN

Orchidaceae

Epipactis gigantea Dougl. ex Hook.

NA

CR

Habenaria elegans (Lindl.) Boland.
Habenaria unalascensis (Spreng.)

NA

RO

CR

CA

Wats.

NA

CR

CA

Poaceae

Agrostis diegoensis Vasey

NA

RO

CR

CA

CL

Agrostis exarata Trin.

Agrostis semiverticillata (Forsk.) C.

NA

RO

CR

CA

Chr.

IN

RO

CR

AN

CA

Agrostis stolonifera L. var. major

(Gaudin) Farw.

IN

85

Ammophila arenaria (L.) Link

IN

NI

Andropogon glomeratus (Walt.) BSP.
Aristida adscensionis L. var. modesta

NA

CR

Hack, in Stuckert

NA

RO

CR

31

CA

CL

GU

Aristida divaricata Humb. & Bonpl. in

Willd.

NA

CR

Arundo donax L.

IN

NI

Avena barbata Brot.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Avena fatua L.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Avena sativa L.

IN

RO

CR

CA

CL

39

Bothriochloa barbinodis (Lag.) Herter

NA

31

CA

Brachypodium distachyon (L.) Beauv.

IN

CA

Bromus arizonicus (Shear) Steb.

NA

Ml

RO

83

AN

NI

BA

CA

CL

Bromus carinatus H. & A.

NA

Ml

RO

CR

AN

NI

CA

CL

Bromus diandrus Roth

IN

Ml

RO

CR

AN

NI

BA

CA

CL

GU

Bromus madritensis L.

IN

CR

26

Bromus maritimus (Piper) Hitchc.

NA

MI

RO

CR

AN

NI

CA

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 31

Taxon

Occur-

rence

San

Miguel

Island

Santa

Rosa

Island

Santa

Cruz

Island

Anacapa

Island

San

Nicolas

Island

Santa

Barbara

Island

Santa San Isla

Catalina Clemente Guada-
Island Island lupe

Bromus mollis L.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Bromus pseudolaevipes Wagnon

NA

RO

CR

CA

Bromus rubens L.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Bromus stamineus Desv. in Gray

IN

CR

Bromus sterilis L.

IN

CA

Bromus tectorum L.

IN

GU

Bromus trinii Desv. in C. Gay

NA

46

RO

CR

AN

NI

BA

CA

CL

GU

Bromus vulgaris (Hook.) Shear

NA

31

5

6

Calamagrostis rubescens Buckl.

NA

CR

Cortaderia atacamensis (Phil.) Pilg.

IN

CR

CA

Crypsis aculeata (L.) Ait.

IN

CA

Cynodon dactylon (L.) Pers.

IN

31

RO

CR

AN

NI

CA

CL

Dactylis glomerata L.

IN

CA

CL

Deschampsia danthonioides (Trin.)

Munro in Benth.

NA

CL

Dissanthelium californicum (Nutt.)

Benth.

EN

CA

CL

GU

Distichlis spicata (L.) Greene var. sto-

lonifera Beetle

NA

MI

RO

CR

AN

NI

CA

CL

Echinochloa crus-galli (L.) Beauv. var.

crus-galli

IN

CA

Ehrharta calycina Sm.

IN

CA

CL

Elymus condensatus Presl

NA

MI

RO

CR

AN

CA

CL

Elymus glaucus Buckl. ssp. glaucus

NA

46

CR

CA

Elymus pacifcus Gould

NA

MI

81

Elymus triticoides Buckl.

NA

MI

RO

CR

AN

CA

Festuca arundinacea Schreb.

IN

CA

Gastridium ventricosum (Gouan)

Schinz & Thell.

IN

RO

CR

CA

CL

Hordeum brachyantherum Nevski

NA

33

Hordeum californicum Covas. & Steb.

NA

MI

RO

CR

AN

NI

CA

Hordeum depressum (Scribn. & Sm.)

Rydb.

NA

CR

33

Hordeum geniculatum All.

IN

RO

CR

CA

CL

Hordeum murinum L. ssp. glaucum

(Steud.) Tzvel.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Hordeum murinum L. ssp. leporinum

(Link) Arcang.

IN

MI

RO

CR

AN

NI

CA

CL

GU

Hordeum pusillum Nutt.

NA

MI

CR

AN

BA

CA

CL

Hordeum vulgare L.

IN

RO

CR

CA

CL

Koeleria pyramidata (Lam.) Beauv.

NA

RO

CR

Lamarckia aurea (L.) Moench

IN

MI

RO

CR

AN

NI

BA

CA

CL

Lolium perenne L. ssp. multiflorum

(Lam.) Husnot

IN

CR

NI

CA

CL

Lolium perenne L. ssp. perenne

IN

CR

NI

CA

CL

Lolium strictum Presl

IN

CA

Lolium temulentum L.

IN

RO

CR

CA

CL

Melica imperfecta Trin.

NA

MI

RO

CR

AN

BA

CA

CL

GU

Monanthochloe litt oralis Engelm.

NA

MI

RO

CR

CA

32 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Taxon

Occur-

rence

San

Miguel

Island

Santa

Rosa

Island

Santa

Cruz

Island

Anacapa

Island

San

Nicolas

Island

Santa

Barbara

Island

Santa

Catalina

Island

San

Clemente

Island

Isla

Guada-

lupe

Muhlenbergia microsperma (DC.)

Kunth

NA

RO

CR

AN

BA

CA

CL

GU

Oryzopsis miliacea (L.) Benth. Hook.

ex Aschers. & Schweinf.

IN

RO

CR

CA

Parapholis incurva (L. f.) C.E. Hubb.

IN

MI

RO

CR

NI

BA

CA

CL

Paspalum dilitatum Poir. in Lam.

IN

CA

Paspalum distichum L.

IN

81

CA

Pennisetum clandestinum Hochst. ex

Chiov.

IN

81

CR

Phalaris aquatica L.

IN

CA

Phalaris canariensis L.

IN

42

41

CA

Phalaris caroliniana Walt.

NA

CR

NI

BA

CA

CL

GU

Phalaris lemmonii Vasey

NA

RO

CA

CL

Phalaris minor Retz.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Phalaris paradoxa L.

IN

CL

Poa annua L.

IN

MI

RO

CR

31

CA

CL

GU

Poa bolanderi Vasey ssp. howellii

(Vasey & Scribn.) Keck

NA

CR

Poa douglasii Nees

NA

MI

RO

Poa palustris L.

NA

CA

Poa scabrella (Thurb.) Benth. ex

Vasey

NA

RO

CR

AN

CA

CL

GU

Polypogon interrupt us HBK.

IN

MI

RO

CR

CA

CL

Polypogon monspeliensis (L.) Desf.

IN

MI

RO

CR

NI

BA

CA

CL

GU

Schismus arabicus Nees

IN

CA

Schismus barbatus Thell.

IN

CR

Scleropoa rigida (L.) Griseb.

IN

CA

Sitanion jubatum J.G. Sm.

IN

CA

Sorghum bicolor (L.) Moench

IN

CA

Sorghum halepense (L.) Pers.

IN

CA

Stipa cernua Steb. & Love

NA

RO

CR

AN

NI

CA

CL

Stipa Columbiana Macoun var. nelsoni

(Scribn.) Hitchc.

NA

NI

Stipa lepida Hitchc.

NA

MI

RO

CR

AN

31

BA

CA

CL

GU

Stipa pulchra Hitchc.

NA

MI

RO

CR

AN

NI

BA

CA

CL

Triticum aestivum L.

IN

CL

Triticum cylindricum (Host.) Ces.,

Pass. & Gib.

IN

CR

Vulpia bromoides (L.) S.F. Gray

IN

MI

RO

CR

AN

31

CA

CL

GU

Vulpia microstachys (Nutt.) Benth.

var. pauciflora (Beal) Lonard &

Gould

NA

MI

5

5

AN

31

CA

CL

GU

Vulpia myuros (L.) K.C. Gmelin var.

hirsuta Hack.

IN

MI

RO

CR

AN

NI

BA

CA

CL

GU

Vulpia myuros (L.) K.C. Gmelin var.

myuros

IN

CA

CL

Vulpia octoflora (Walt.) Rydb. var.

hirtella (Piper) Henr.

NA

MI

RO

CR

AN

NI

BA

CA

CL

GU

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 33

Taxon

Occur-

rence

San

Miguel

Island

Santa

Rosa

Island

Santa

Cruz

Island

Anacapa

Island

San

Nicolas

Island

Santa

Barbara

Island

Santa San Isla

Catalina Clemente Guada-
Island Island lupe

Potamogetonaceae

Potamogeton crispus L.

IN

CA

Potamogeton pectinatus L.

NA

RO

CR

CA

Ruppia maritima L.

NA

RO

CR

CA

CL

96

Typhaceae

Typha domingensis Pers.

NA

MI

RO

CR

NI

CA

Typha latifolia L.

NA

NI

CA

CL

Zosteraceae

Phyllospadix scouleri Hook.

NA

66

AN

NI

BA

CA

CL

Phyllospadix torreyi Wats.

NA

MI

RO

CR

AN

NI

BA

CA

CL

GU

Zostera marina L.

NA

RO

CR

AN

CA

GU

Note 1. Davis (1980) proposed these notations for new taxa he recognizes as a result of his detailed studies of Malacothrix on the Channel
Islands. They are recognized here since placement into earlier associated taxa would add to the systematic confusion. Dr. Davis is preparing
a manuscript treating these taxa (pers. comm.).

Note 2. Morin (Systematic Botany 8(4):436— 468. 1983) separates the material from Guadalupe Island to var. guadalupensis Morin.

Note 3. A detailed discussion by Philbrick (1980) indicated that this species was native on MI, AN, CA, CL, and possibly native on RO, and
NI, but probably cultivated on CR. See also Eastwood (1941).

Note 4. Probably introduced on CR (RSA-POM; SBBG).

Note 5. Probably introduced on CA (Moran 596 LAM!).

FLORISTIC RELATIONSHIPS

The following discussion concerns the distribution of native
vascular plant taxa on the islands and the floristic relation-
ships among the various islands. Analysis of dispersal types
and patterns as well as evolutionary histories for each taxon
would be needed to provide information on the origins of
the insular floras and is not in the scope of this work. Intro-
duced taxa are generally distributed among the islands and
will not be discussed here further (Table 1). Map 1 shows
the spacial relationships among the islands while Table 1 and
Table 2 summarize the distributions of the taxa.

Nearly 275 of the taxa native to the islands are found on
one or more of the islands of the northern group (San Miguel,
Santa Rosa, Santa Cruz, and Anacapa), one or more of the
islands of the southern group (San Nicolas, Santa Barbara,
Santa Catalina, and San Clemente), and are generally dis-
tributed on adjacent mainland areas. Some of these taxa are
also native to Guadalupe Island, Mexico.

The northern islands exhibit close floristic affinities. The
percentages of native taxa shared among the islands are high.
The numbers of shared insular endemics are also high. Thir-
ty-eight of the 58 shared insular endemics occur on the north-
ern islands.

Several groupings of native taxa are worthy of note. The
first is native taxa found on the northern islands, generally
on the mainland, but absent from the southern islands. The
second group consists of taxa found on the southern islands,
generally on the mainland, but absent from the northern

islands. The third group is composed of taxa which have the
southern limits of their ranges on the islands or adjacent
mainland areas. The fourth group contains taxa with the
northern limits of their ranges on the islands or adjacent
mainland areas. There are 39 taxa with mainland ranges
which terminate considerably north of their insular occur-
rences. Twenty-three taxa have mainland ranges which ter-
minate considerably south of their insular occurrence. There
are also a few taxa which seem to have insular occurrences
which represent westerly extensions of inland ranges.

Taxa which occur on one or more of the islands of the
northern group but are absent from the southern islands even
though their mainland ranges extend far to the south include
the following:

Cystopteris fragilis

Polystichum munitum ssp. munitum

Woodwardia fimbriolata

Cheilanthes clevelandii

Pinus torreyana

Acer macrophyllum

Benda erecta

Lomatium utriculatum

Agoseria grandi flora

Aster exilis

Chaenactis glabriuscula var. lanosa
Gnaphalium ramosissimum
Lasthenia glabrata ssp. coulteri

34 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Layia glandulosa ssp. glandulosa

Layia platyglossa ssp. platyglossa

Lepidospartum squamatum

Micropus californieus

Stephanomeria cichoriacea

Venegasia carpesioides

Cryptantha muricata var. jonesii

Pectocarya penicillata

Plagiobothrys californieus var. fulvescens

Erysimum ammophilum

Thysanocarpus curvipes var. curvipes

Thysanocarpus laciniatus var. crenatus

Cardionema ramosissima

Cuscuta ceanothi

Cuscuta salina

Dudleya caespitosum

Cucurbit a foetidissima

Arbutus menziesii

Vaccinium ovatum

Lotus micranthus

Lupinus bicolor ssp. tridentatus

Lupinus polycarpus

Trifolium variegatum

Vicia americana

Quercus agrifolia

Quercus wislizenii var. frutescens

Centaurium exaltatum

Phacelia ramosissima var. austrolittoralis

Salvia brandegei

Stachys bullata

Lythrum californicum

Camissonia hirtella

Camissonia ignota

Camissonia strigulosa

Ludwigia peploides

Oenothera elata ssp. hirsutissima

Oxalis albicans ssp. pilosa

Meconella denticulata

Papaver californicum

Plantago bigelovii ssp. californica

Plantago hirtella var. galeottiana

Plantago maritima

Claytonia perfoliata var. parviflora

Montia fontana ssp. amporitana

Anagallis minima

Rumex fueginus

Samolus parviflora

Clematis lasiantha

Ranunculus californieus var. californieus

Rhamnus californica ssp. californica

Galium californicum ssp. flaccidum

Lithophragma cymbalaria

Saxifraga californica

Diplacus longiflorus

Mimulus nasutus

Petunia parviflora

Carex barbarae

Carex globosa

Carex senta

Scirpus californieus

Scirpus cernuus ssp. californieus

Juncus efl'usus var. pacificus

Juncus phaeocephalus

Lilium humboldtii ssp. ocellatum

Zigadenus fremontii var. fremontii

Epipactis gigantea

Aristida divaricata

Hordeum depressum

Poa bolanderi ssp. howe/lii

Several taxa likewise occur on one or more islands of the
southern group but are not known from the northern islands
even though their mainland ranges may extend far to the
north. These are as follows:

Azol/a filiculoides
Malosma laurina

Ambrosia psilostachya var. californica
Artemisia dracunculus
Baccharis emoryi

Haplopappus pa/meri ssp. pachylepis
Helenium puberulum
Malacothrix saxatilis var. tenuifolia
Microseris douglasii ssp. douglasii
Pluchea sericea

Xanthium strumarium var. canadense

Cryptantha intermedia

Cryptantha microstachys

Lepidium virginicum var. pubescens

Thysanocarpus curvipes var. elegans

Tropidocarpum gracile

Callitriche marginata var. marginata

Callitriche longipedunculata

Atriplex serenana var. serenana

Bassia hyssopifolia

Convolvulus si mu I a ns

Cuscuta occidentalis

Cornus glabrata

Crassula aquatica

Elatine californica

Euphorbia crenulata

Euphorbia serpyllifolia var. serpyllifolia

Euphorbia spathulata

Nemophila menziesii ssp. menziesii

Juglans californica

Salvia apiana var. apiana

Trichostema lanceolatum

Eremalche exilis

Malvella leprosa

Camissonia cheiranthifolia ssp. suffruticosa
Allophyllum glutinosum
Navarretia hamata ssp. foliosa
Eriogonum fasciculatum ssp. fasciculatum
Potentilla glandulosa ssp. glandulosa
Galium angustifolium ssp. angustifolium
Lithophragma affine ssp. mixtum
Castilleja foliolosa

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 35

Diplacus puniceus

Mimulus brevipes

Verbena bracteata

Vitis girdiana

Scirpus microcarpus

Juncus acutus var. sphaerocarpus

Juncus textilis

Calochortus splendens

Deschampsia danthonioides

Stipa columbiana var. nelsonii

Typha latifolia

The ranges of the following taxa have their southern limits
on the islands or nearby mainland areas:

Equisetum hyema/e ssp. affine

Athyrium felix-femina var. sitchensis

Adiantum pedatum ssp. aleuticum

Pinus muricata

Lomatium caruifolium

Agoseris apargioides ssp. apargioides

Aster chilensis

Aster radulinus

Baccharis plummerae

Blennosperma nanum

Cirsium brevistylum

Cirsium proteanum

Erigeron glaucus

Erigeron sanctarum

Eriophyllum staechadifolium var. artemisiaefolium

Eriophyllum staechadifolium var. depressum

Grindelia latifolia

Grindelia stricta ssp. venulosa

Haplopappus squarrosus ssp. grindelioides

Haplopappus venetus var. sedoides

Hemizonia increscens ssp. increscens

Hieraceum argutum

Malacothrix coulteri var. cognata

Microseris douglasii ssp. tenella

Pentachaeta lyonii

Stephanomeria exigua ssp. coronaria

Cryptantha leiocarpa

Erysimum insulare

Silene laciniata ssp. major

Calystegia macrostegia ssp. cyclostegia

Astragalus curvipes

Lotus grandiflorus var. grandiflorus

Lupinus arboreus

Lupinus densiflorus var. palustris

Pickeringia montana

Trifolium barbigerum

Garrya elliptica

Erodium macrophyllum var. californicum
Phacelia ramosissima var. montereyensis
Lepechinia fragrans
Satureja douglasii
Hesperolinon micranthum

Ammannia coccinea

Malacothamnus fasciculatus ssp. catalinensis
Sidalcea malvaeflora ssp. malvaeflora
Abronia latifolia

Orobanche uniflora ssp. occidentalis

Eschscholzia californica var. maritima

Plantago maritima

Linanthus androsaceus ssp. luteus

Linanthus bicolor ssp. bicolor

Chorizanthe wheeleri

Calandrinia breweri

Dodecatheon clevelandii ssp. insularis

Ranunculus californicus var. cuneatus

Holodiscus discolor ssp. discolor

Potentilla egedii var. grandis

Salix lasiandra var. lasiandra

Ribes malvaceum

Ribes menziesii var. menziesii

Antirrhinum multiflorum

Mimulus cardinalis

Mimulus floribundus

Mimulus guttatus ssp. littoralis

Mimulus guttatus ssp. micranthus

Scrophularia californica ssp. californica

Nicotiana bigelovii

Carex gracilior

Carex montereyensis

Carex pansa

Carex subbracteata

Luzula subsessilis

Juncus effusus var. brunneus

Allium lacunosum

Calamagrostis rubescens

Koeleria pyramidata

Poa douglasii

The following taxa have the northern limits of their ranges
on the islands or nearby mainland:

Notholaena californica
Notholaena newberryi
Rhus ovata
Filago arizonica

Microseris douglasii ssp. platycarpha

Stephanomeria diegensis

Lepidium virginicum var. robinsonii

Thysanocarpus laciniatus var. laciniatus

Bergerocactus emoryi

Opuntia oricola

Opuntia prolifera

Cleome isomeris

Aphanisma blitoides

At rip lex coulteri

Atriplex pacifica

Atriplex watsonii

Xylococcus bicolor

Euphorbia miser a

36 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Lathyrus laetiflorus ssp. alefeldii

Lotus heermanmi

Lupinus agardhianus

Phacelia grandiflora

Lepechinia fragrans

Oxalis albicans ssp. californicus

Linanthus dianthiflorus ssp. dianthiflorus

Eriogonum cinereum

Oligomeris linifolia

Ceanothus megacarpus ssp. megacarpus
Antirrhinum nuttallianum
Orthocarpus purpurascens var. pallidus
Lycium californicum
Nicotiana clevelandii
Allium praecox

Lilium humboldtii ssp. ocellatum
Monanthochloe lift oralis

The following taxa have mainland ranges which terminate
considerably north of their insular occurrences:

Polypodium scouleri
Notholaena newberryi
Juniperus californica
Agoseris heterophylla
Coreopsis gigantea

Eriophyllum lanatum var. grandiflorum
Gnaphalium purpureum
Lasthenia coronaria
Malacothrix clevelandii
Malacothrix incana
Pectocarya linearis ssp. ferocula
Pectocarya recurvata

Plagiobothrys californicus var. californicus
Dithyrea maritima

Lepidium lasiocarpum var. latifolium
Stellaria nitens

Lupinus albifrons var. albifrons
Lupinus albifrons var. douglasii
Centaurium davyi
Lepechinia calycina

Camissonia cheiranthifolia ssp. cheiranthifolia

Clarkia davyi

Clarkia prostrata

Epilobium foliosum

Armeria maritima

Allophyllum gilioides

Polygala californica

Myosurus minimus var. filifolius

Ceanothus crassifolius

Cercocarpus betuloides ssp. blancheae

Ribes sanguineum

Castilleja mollis

Orthocarpus attenuatus

Orthocarpus densiflorus ssp. densiflorus

Carex tumulicola

Calochortus luteus

Bromus maritimus
Elymus pacificus

The following taxa have mainland ranges which terminate
considerably south of their insular ranges:

Pityrogramma triangidaris var. viscosa
Pinus torreyana

Haplopappus venetus ssp. furfuraceus
Malacothrix similis
Senecio lyonii
Stephanomeria diegensis
Harpagone/la pa/meri
Plagiobothrys californicus var. gracilis
A triplex pacifica

Astragalus trichopodus ssp. lecopsis

Lotus scoparius var. veatchii

Quercus douglasii

Pholistoma racemosum

Salvia brandegei

Mentzelia affinis

Camissonia robusta

Eriogonum grande ssp. grande

Calandrinia maritima

Prunus lyonii

Ribes viburnifolium

Galvezia speciosa

Lycium fremontii

Insular occurrences for a few taxa seem to be extensions
of generally interior distributions. These include:

Sarcostemma cynanchoides
Thelesperma megapotamicum
Cryptantha maritima
Mono/epis nuttalliana
Quercus engelmannii
Erodium texanum
Populus fremontii ssp. fremontii
Lycium brevipes var. brevipes
Andropogon glomeratus
Poa palustris

The islands of the northern group exhibit closer floristic
similarities to one another than do the islands of the southern
group. The islands lie close to one another as well as the
mainland (Table 1 and Table 2).

Only five taxa, including one endemic, occur on San Miguel
Island to the exclusion of the other islands. San Miguel shares
90% of its shared native taxa with Santa Rosa Island and
89% with Santa Cruz Island. San Miguel and Santa Rosa also
share alone eight taxa with the mainland and two insular
endemics. Though separated from each other, San Miguel
and Santa Cruz share alone four taxa with the mainland and
one insular endemic. The floristic complement found on San
Miguel Island has similarities with that of Santa Rosa Island
as well as Santa Cruz Island.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 37

Map 1. Channel Islands of Southern California and Isla Guadalupe, Mexico.

Santa Rosa Island is close to the mainland, and Santa Cruz
Island. There are 27 nonendemic native taxa found exclu-
sively on Santa Rosa and the mainland. There are only three
unshared endemic taxa on the island, despite its large size,
perhaps indicative of less isolation than some of the other
islands. There are seven endemics shared exclusively with
Santa Cruz Island. There are 3 1 native taxa restricted to Santa
Rosa, Santa Cruz and the mainland. Santa Rosa shares 92%
of its shared native taxa with Santa Cruz. The floristic affin-
ities of Santa Rosa Island lie with Santa Cruz Island.

Santa Cruz Island is the largest of the Channel Islands of

southern California. Concomitant with its size, ecological
diversity, and proximity to the mainland, there are 6 1 native
taxa found exclusively on Santa Cruz and the mainland. This
is comparable to 58 such taxa found on Santa Catalina Island
to the south. Both islands are at similar distances from the
mainland. The fact that 1 7 of the 1 9 taxa endemic to more
than one of the northern islands occur on Santa Cruz is
indicative of the close affinities of Santa Cruz to the other
islands of the northern group. A total of 16 taxa are found
on the southern islands and Santa Cruz alone of the northern
islands.

38 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Table 1. Features of the Channel Islands of southern California and Isia Guadalupe, Mexico and floristic data for each. Area, and highest
altitude, as well as shortest distance to the mainland are given for each island. The numbers of native and introduced taxa are given as are
numbers of unshared endemic taxa for each island. The number of insular endemics shared with the other islands dealt with here are also
listed for each of the islands.

Island

Area in km2

Highest
altitude in
meters

Distance to
mainland in km

Number of
native taxa

Number of
introduced
taxa

Number of
unshared
insular
endemics

Number of
shared
insular
endemics

San Miguel

36

262

42

171

50

1

10

Santa Rosa

218

480

44

370

80

3

35

Santa Cruz

244

660

31

477

137

10

35

Anacapa

3

284

21

166

40

1

18

San Nicolas

57

277

98

1 14

66

4

13

Santa Barbara

3

193

61

72

29

3

8

Santa Catalina

194

631

32

417

175

8

24

San Clemente

145

589

79

259

83

14

29

Guadalupe

249

1298

253

168

38

35

16

Some of these data are from Philbrick (1967).

Anacapa is a series of small islands forming an eastward
extension of Santa Cruz Island. There are only two taxa,
including one endemic, that occur on Anacapa Island to
the exclusion of the other islands. There are many more taxa
on Anacapa than on the more isolated Santa Barbara Island
which is of similar size. Anacapa shares 95% of its shared
native taxa with Santa Cruz Island and 89% with Santa Rosa
Island.

The islands of the southern group are more widely scattered
than those of the northern group. The floras are generally
more dissimilar among the islands.

San Nicolas Island has two endemic taxa and an additional
four taxa restricted to San Nicolas and the mainland. San
Nicolas shares about 77% of its shared native taxa with Santa
Cruz Island and the same percentage with Santa Catalina
Island, although the individual species shared may be dif-
ferent. San Nicolas shares three native taxa exclusively with
Santa Cruz and the mainland and only two native taxa ex-
clusively with Santa Catalina and the mainland. San Nicolas
shares eight insular endemics exclusively with the southern
islands and only one exclusively with the northern islands.
The affinities of San Nicolas Island lie equally with the north-
ern and southern islands but the balance seems to favor
relationship to the southern islands in light of the high num-
ber of shared insular endemics.

Only the three endemic taxa of the 72 native taxa on Santa
Barbara Island are not shared by another island. Santa Bar-
bara shares 89% of its shared native taxa with Santa Catalina
Island and San Clemente Island and 81% with Santa Cruz
Island. Santa Barbara shares six of the endemic taxa exclusive
to the southern group of islands.

Santa Catalina Island is the largest island of the southern
group. As noted above, there are 58 native taxa exclusive to
Santa Catalina and the mainland. Nine of the 16 taxa en-
demic to more than one island of the southern group are
found on Santa Catalina. Santa Catalina shares 86% of its
shared native taxa with Santa Cruz Island. There are 37

native taxa common only to Santa Catalina, Santa Cruz, and
the mainland. Five taxa are endemic to the northern islands
and only Santa Catalina of the southern group. A total of 65
taxa are native to the northern islands and only Santa Cat-
alina Island of the southern group, as well as the mainland.
The floristic affinities of Santa Catalina Island lie with the
mainland and the Santa Cruz Island area.

There are 14 taxa endemic to San Clemente Island and
only five taxa restricted to San Clemente and the mainland.
Ten native taxa are exclusively shared by San Clemente and
Santa Catalina of the islands, and the mainland. San Cle-
mente shares 87% of its shared native taxa with Santa Cat-
alina and 77% with Santa Cruz Island. San Clemente and
Santa Catalina share exclusively three endemic taxa. Twelve
taxa are endemic to San Clemente and one or more of the
other islands of the southern group. Three endemics are com-
mon to the northern islands and San Clemente Island alone
of the southern islands. One insular endemic is shared with
Islas Los Coronados and three are shared with Guadalupe
Island. The floristic affinities of San Clemente Island seem
to lie with Santa Catalina Island.

Guadalupe Island is the largest and most isolated of the
islands considered here. There are 35 taxa endemic to the
island. Twenty-five native taxa are found on Guadalupe and
the mainland but are absent from the Channel Islands. Gua-
dalupe shares 88% of its shared native taxa with Santa Cat-
alina, 79% with San Clemente, and 76% with Santa Cruz.
Guadalupe Island shares ten endemic taxa with one or more
of the Channel Islands. One endemic is shared with Santa
Catalina, four nonendemic native taxa are also shared ex-
clusively between these islands and the mainland. Guadalupe
shares four endemics with San Clemente and two nonen-
demics with San Clemente and the mainland only. Guada-
lupe shares five of the 14 taxa endemic to more than one
island of the southern group only. There are clear floristic
affinities between Guadalupe Island and the Channel Islands
of southern California.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 39

Table 2. Shared native and endemic taxa between island pairs. Number of native taxa, including endemics, shared between island pairs is
found in the coordinate box between any two islands in question above the diagonal of dashes. Similarly number of insular endemics shared
among any two islands is found below the diagonal of dashes.

Total number of shared native taxa (including endemics) between island pairs

San

Miguel

Santa

Rosa

Santa

Cruz

Anacapa

San

Nicolas

Santa

Barbara

Santa

Catalina

San

Clemente

Guadalupe

Number of shared island endemics
between island pairs
San Miguel —

150

148

96

69

40

115

91

43

Santa Rosa

10

-

314

147

81

51

237

161

74

Santa Cruz

8

33

—

156

84

56

304

185

82

Anacapa

6

15

16

-

64

54

134

113

54

San Nicolas

3

4

3

3

—

46

84

81

38

Santa Barbara

0

2

2

2

7

—

62

62

40

Santa Catalina

3

14

14

6

7

6

—

209

96

San Clemente

3

1 1

10

6

10

8

17

—

86

Guadalupe

1

6

6

3

5

4

11

14

—

Many elements of the floras of the Channel Islands of
southern California and Guadalupe Island, Mexico, represent
surviving populations of mainland taxa. Decreased mainland
ranges occasionally create great disjunctions. There are more
taxa from northerly distributions surviving on the islands
than taxa from predominately southern ranges. This is evi-
dent in the above list of taxa which occur on the northern
islands, generally on the mainland, but skip the southern
islands and the list of taxa with mainland ranges which ter-
minate considerably north of their insular occurrences. Shared
insular endemics would seem to indicate stronger floristic
affinities between islands than sharing of other native taxa.

San Miguel and Anacapa islands are not very distinctive
in their floristic composition from that of Santa Rosa and
Santa Cruz islands. The portions of the floras of San Miguel
and Anacapa islands that are shared with the mainland or
other islands are high. Santa Rosa Island exhibits some flo-
ristic divergence from Santa Cruz Island, but is still quite
close in its floristic affinities with Santa Cruz. The northern
group of islands share high percentages of their native taxa,
most have relatively low numbers of unshared endemics, and
more shared insular endemics. This seems to portray a more
unified floristic region.

The floristic affinities among the southern group of islands
are not as pronounced (Table 2). San Nicolas has similar
floristic affinities overall with Santa Cruz and Santa Catalina
islands but shares more insular endemics with the southern
group of islands. Santa Barbara Island shares a higher per-
centage of its flora than does San Nicolas. Santa Barbara
shares a greater percentage of its total native taxa as well as
those included which are insular endemics with islands of
the southern group. Santa Catalina Island has significant flo-
ristic affinity with Santa Cruz Island of the northern group
but shares fewer of the endemics restricted to more than one
of the southern islands. San Clemente has the most unique

floristic elements of the southern islands. It exhibits closest
floristic similarity to Santa Catalina Island. All of the islands
of the southern group, except Santa Catalina, seem to show
greater floristic similarity to other islands of the group than
to mainland areas. The greater distances among these islands
and to mainland areas, as well as lesser floristic similarities
among the islands, indicate that the southern islands are a
less cohesive floristic region than the northern islands.

Guadalupe Island, Mexico, has the most unique floristic
complement of any of the islands treated here. The relation-
ship of the flora of the island to the Channel Islands of south-
ern California is, however, significant.

APPENDIXES

APPENDIX I. SELECTED EXSICCATAE AND
HERBARIA HOUSING INSULAR COLLECTIONS

The citation of herbaria and specimens here provides doc-
umentation of the plants of the Channel Islands and Gua-
dalupe Island. Primary consideration is given to the signif-
icant insular collections combined at LAM. Most certainly
numerous additional specimens are available at RSA-POM
and SBBG, institutions considered among the primary cen-
ters of insular collections. Specimens from these institutions
are cited here as well as from several other herbaria.

The taxa are arranged in the same systematic order found
in the table of vascular plants. The islands also are recorded
in the same sequence. Each collection citation is followed by
the one or more standard herbarium acronyms as given by
Holmgren et al. (1981); any not listed in this publication are
written out more fully. An exclamation (!) following the ac-
ronym indicates collections verified by the author. Acronyms
separated by commas indicate duplicate sheets at other in-
stitutions. No more than three collections are given for a
particular insular record. Herbarium acronyms not following

40 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

a collection citation and lacking an exclamation (!), indicate
herbaria containing specimens which were not specifically
verified. Following all three of the collection citations ad-
ditional herbarium acronyms not accompanied by excla-
mations, indicate herbaria containing additional collections
documenting the insular record. Citations and locations of
additional verified specimens will be offered to interested
monographers, where possible.

In a very few cases an insular record is given as a literature
reference to a particular specimen which I have not yet seen
or been able to locate. These are merely for the reader’s
information, although those considered most reliable were
used in the numerical tabulations of the flora.

Vascular Cryptogams

Selaginellaceae
Selaginella bigelovii Underw.

Santa Rosa: Thorne et al. 48854 RSA!; SBBG; SBM.
Santa Cruz: Clokey 4849 LAM!; Raven & Smith 15270
RSA!; Abrams & Wiggins 341 DS!; SBM.

Anacapa: SBBG.

Santa Catalina: Trask s.n. in Mar. 1900 LAM!; Fosberg
R96 LAM!; Dunkle 1927 LAM!; CAS; DS; RSA-POM.
San Clemente: Dunkle 7228 LAM!; Elmore 398 AHFH!;
Thome 36120 RSA!; CAS.

Equisetaceae

Equisetum hyemale L. spp. affine (Engelm.) Calder & Taylor
Santa Cruz: Hoffmann s.n. Sep. 20, 1930 POM!; SBM.
Equisetum laevigatum A. Br.

Santa Rosa: Thorne et al. 48740 RSA!.

Santa Cruz: Hoffmann s.n. Jul. 1, 1930 LAM!; Raven &
Smith 15269 RSA!; Howell 6271 CAS!; SBBG; SBM.
Santa Catalina: Dunkle 1964 LAM!; Fosberg R90 LAM!;
Thome & Everett 34959 RSA!; SBM.

Equisetum telmateia Ehrh. var. braunii Milde

Santa Cruz: Clokey 5162 LA!, RSA!; Miller s.n. Jun. 6,
1918 CAS#93549!; SBM.

Santa Catalina: Trask s.n. in Jul. 1 900 LAM!; Dunkle 2458
AHFH!; Wolf 3607 RSA!; SBM.

Aspidiaceae

Athyrium felix-femina (L.) Roth var. sitchensis Rupr.

Santa Cruz: Clokey 4844 POM!; Munz & Crow 11833
POM!; Wolf 2915 Mar. 27, 1932 RSA!; SBBG; SBM.
Cyst opt eris fragilis ( L. ) Bern h .

Santa Cruz: Hoffmann s.n. May 23, 1932 POM!; Hoff-
mann s.n. May 23, 1932 CAS#194980!; SBM.
Dryopteris arguta (Kaulf.) Watt
Santa Rosa: Dunkle 8429 LAM!; Thome et al. 48836 RSA!;

Munz & Crow 1 1645 POM!; SBM.

Santa Cruz: Dunkle 8602 LAM!, DS!; Clokey 4844 LAM!;

Blakley 3379 RSA!; CAS; SBM.

Anacapa: Blakley 5765 SBBG!.

Santa Catalina: Dunkle 1948 LAM!; Fosberg R85 LAM!;

Thome & Everett 34607 RSA!; DS; SBM.

San Clemente: Munz 6706 POM!.

Polystichum munitum (Kaulf.) Presl ssp. munitum

Santa Rosa: SBM.

Santa Cruz: Pierson 11083 RSA!; Munz & Crow 11887
POM!; SBBG; SBM.

Polystichum munitum (Kaulf.) Presl ssp. solitarium Maxon

Guadalupe: Anthony 9 NY!; Palmer 102 NY!.

Blechnaceae

Woodwardia fmbriata Sm. in Rees

Santa Rosa: SBM.

Santa Cruz: Clokey 4842 LAM!; Dunkle 8604 LAM!; Balls
& Blakley 23741 RSA!; SBM.

Polypodiaceae

Polypodium californicum Kaulf.

Santa Rosa: Thorne et al. 48817 RSA!; SBM.

Santa Cruz: Dunkle 8620 LAM!; SBM.

Anacapa: SBBG; SBM.

Santa Barbara: SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Dunkle
2044 LAM!; Dunkle 2045 LAM!; RSA-POM.

San Clemente: RSA-POM.

Guadalupe: Franceschi s.n. US!; Brandegee s.n. Mar. 23,
1897 US!; Palmer 857 US!.

Polypodium scouleri Hook. & Grev.

Santa Cruz: RSA-POM; SBM.

Guadalupe: Palmer 104 NY!; Moran 6422 RSA!.

Pteridaceae

Adiantum capillus-veneris L.

Santa Rosa: Thome et al. 48800a RSA!; SBBG; SBM.

Santa Cruz: Pierson 1 1089 RSA!; Balls & Blakley 23724
RSA!; Williams 42 POM!; SBM.

Anacapa: Hoffmann s.n. May 16, 1929 SBM!; Bond 347
SBM!; SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Fosberg
R94 LAM!; Thome & Everett 35001 RSA!.

Adiantum jordani C. Muell.

Santa Rosa: Thome et al. 48990 RSA!; Blakley 3097 RSA!;
Dunn, D. 1335 LA!; SBM.

Santa Cruz: Dunkle 8537 LAM!; Clokey 4831 LAM!; Ra-
ven & Smith 15177 RSA!; SBM.

Anacapa: SBBG; SBM.

Santa Catalina: Grant s.n. Apr. 20, 1904 LAM!; Fosberg
R79 LAM!; Thome & Everett 34486 RSA!; SBM.

San Clemente: House & Grumbles s.n. Aug. 5, 1913 USC!;
Thome 42967 RSA!; Raven 17694 RSA!; SBBG; DS.

Adiantum pedatum L. ssp. aleuticum (Rupr.) Calder & Taylor

Santa Cruz: Clokey 4828 LAM!; Munz & Crow 11839
POM!; Wolf 2912 RSA!; SBBG; SBM.

Aspidotis californica (Hook.) Nutt, ex Copel.

Santa Cruz: Hoffmann 1 73 SBM!; Clokey 5 157 GH!; Munz
& Crow 11852 POM!.

Santa Catalina: Trask s.n. in Mar. 1 897 LAM!, MO!; Dun-
kle 1879 AHFH!; Thome & Everett 34848 RSA!.

Cheilanthes clevelandii D.C. Eat.

Santa Rosa: Dunkle 8512 AHFH!; Spencer 199 US!; Bran-
degee s.n. in 1888 UC!.

Santa Cruz: Clokey 4838 LAM!; Clokey 5158 LA!, RSA!,
UC!; Brandegee s.n. in Apr. 1888 SBM# 15936!, UC!.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 41

Cheilanthes newberryi (D.C. Eat.) Domin
San Clemente: Dunkle 7230 LAM!; Trask 326 NY!; DeBuhr
& Wallace 709 RSA!.

Guadalupe: Franceschi 39 LAM!, MO!; Moran 18147 SD!;
Palmer 105 MO!.

Notholaena californica D.C. Eat.

Santa Catalina: Trask s.n. in 1899 MO!; Blakley 5510
SBBG!.

Guadalupe: Moran 6165 DS!, SD!.

Pellaea andromedaefolia (Kaulf.) Fee var. pubescens D.C.
Eat.

Santa Rosa: Dunn, N. s.n. LA!; Thome 48865 RSA!; Ra-
ven, Blakley & Omduff 14968a RSA!; SBM.

Santa Cruz: Wolf 2778 RSA!; Blakley 3301 RSA!; Raven
& Smith 15258 RSA!; SBM.

Anacapa: Dunkle 7663 LAM!; SBM.

Santa Catalina: Trask s.n. in Mar. 1900 LAM!; Fosberg
R86 LAM!; Wolf 3450 RSA!.

San Clemente: Dunkle 7247 LAM!; Elmore 413 AHFH!;
Thome 42867 RSA!.

Pellaea mucronata (D.C. Eat.) D.C. Eat. ssp. mucronata
Santa Rosa: Epling & Erickson s.n. Aug. 8, 1937 LA!;
Hoffmann 154 POM!; SBM.

Santa Cruz: Clokey 4840 LAM!, NY!, POM!; Hoffmann
s.n. LAM!; SBM.

Santa Catalina: Trask s.n. in Jan. 1896 US!; Fosberg R84
LAM!, NY!, US!; Dunkle 1872 AHFH!; Thome & Ev-
erett 34991 RSA!.

Guadalupe: Palmer 100 NY!; Moran 6166 RSA!, SD!;
Moran 7838 RSA!, SD!.

Pityrogramma triangularis (Kaulf.) Maxon var. triangularis
Santa Rosa: Thome et al. 48900 RSA!; RSA-POM; SBBG;
SBM.

Santa Cruz: SBBG; SBM.

Anacapa: Dunkle 7664 AHFH!; SBBG; SBM.

Santa Catalina: RSA-POM; SBM.

San Clemente: Dunkle 7244 NY!; Thome 42866 RSA!.
Guadalupe: Mason 1514 CAS!; Moran 7833 DS!, SD!;
Palmer 856 NY!, US!.

Pityrogramma triangularis (Kaulf.) Maxon var. viscosa (D.C.
Eat.) Weath.

Santa Rosa: Hoffmann s.n. Mar. 26, 1927 SBM#507!.
Santa Cruz: Munz & Crow 1 1544 LA!; Clokey 4824 NY!;
Clokey 5152 NY!; SBM.

San Nicolas: Newman 124 Pt. Mugu Nav. Air. Sta.!.
Santa Catalina: Trask s.n. in Mar. 1901 NY!; Rose, Georgia
s.n. in 1889 NY!; SBBG.

San Clemente: Trask 328 NY!; Trask 329 NY!; SBBG.
Pteridium aquilinum (L.) Kuhn var. pubescens Underw.
Santa Rosa: Dunkle 8511 LAM!, AHFH!, RSA!; Blakley
3177 RSA!; Munz & Crow 1 1668 POM!; SBM.

Santa Cruz: Dunkle 8603 LAM!, AHFH!, RSA!; Clokey
4835 LAM!; Fosberg 7595 LAM!; SBM.

Santa Catalina: Trask s.n. in Aug. 1902 LAM!; Fosberg
S4834 LAM!; Thome & Thome 36429 RSA!; SBM.

Salviniaceae

Azolla filiculoides Lam.

Santa Catalina: Thome 36659 RSA!; Raven 17853 RSA!;
Piehl 62522 RSA!; SBBG.

Gymnosperms

Cupressaceae

Cupressus guadalupensis Wats. ssp. guadalupensis

Guadalupe: Rempel 758-37 LAM!; Ziezenhenne s.n. in
May 1933 AHFH!; Franceschi 3 RSA!; SD.

Cupressus macrocarpa Hartw. ex Gord.

Santa Cruz: SBBG.

Anacapa: SBBG.

Santa Catalina: Parratt 524 LAM!.

Juniperus californica Carr.

Guadalupe: Moran 6454A SD!; Moran 2635A SD!.

Pinaceae

Pinus muricata D. Don

Santa Cruz: Clokey 4850 LAM!, RSA!, POM!; Dunkle
8655 LAM!; Wolf 2844 RSA!.

Pinus radiata D. Don var. binata (Engelm. in Wats.) Lem-
mon

Guadalupe: Palmer 90 NY!; Rose 16001 NY!; Howell 8183
RSA!, NY!; CAS.

Pinus remorata Mason

Santa Rosa: Moran 3348 LAM!; Dunkle 8505 LAM!,
AHFH!; Thome et al. 48983 RSA!; SBM.

Santa Cruz: Clokey 4853 LAM!, POM!; Williams 79 POM!;
Raven & Smith 15317 RSA!; SBM.

Pinus torreyana Parry ex Carr.

Santa Rosa: Dunkle 8500 LAM!, AHFH!, RSA!; Thome
et al. 48852 RSA!; Abrams & Wiggins 247 RSA!.

Dicotyledons

Aceraceae

Acer macrophyllum Pursh

Santa Cruz: Hoffmann s.n. RSA#5771!; Clokey 4995 POM!;
Munz & Crow 1 1868 POM!; SBBG.

Aizoaceae

Aptenia cordifolia (L. f.) N.E. Br.

Anacapa: SBBG.

Santa Catalina: Thome 36726 RSA!; Thome & Everett
33454 RSA!; SBBG.

Carpobrotus aequilaterus (Haw.) N.E. Br.

San Miguel: Dunkle 8413 LAM!, RSA!; Elmore 316
AHFH!; Munz & Norris 11826 POM!; SBBG; SBM.
Santa Rosa: Thome et al. 48918 RSA!; SBBG; SBM.
Santa Cruz: Sauer & Laughrin 5519 RSA!; SBM.
Anacapa: SBBG; SBM.

San Nicolas: SBBG.

San Clemente: Raven 18045 RSA!.

Carpobrotus edulis (L.) Bolus
San Nicolas: RSA; SBBG.

42 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Malephora crocea (Jacq.) Schwant.

Anacapa: Blakley 28 1 3 SBBG!; Benedict s.n. Apr. 22, 1 970
SBBG!.

Mesembryanthemum crystallinum L.

San Miguel: Kanakoff s.n. Apr. 13, 1940 RSA!; SBBG.
Santa Rosa: Dunn, D. 1 346 LA!; Thome et al. 48920 RSA!;

Raven, Blakley & Omduff 14966 RSA!; SBBG; SBM.
Santa Cruz: Ferren 1894 SCIR!; SBBG; SBM.

Anacapa: Dunn, N. s.n. May 14, 1932 LA!; SBBG; SBM.
San Nicolas: Dunkle 8303 LAM!; Raven & Thompson
20704 LA!, RSA!; Foreman & Lloyd 147 RSA!; SBBG;
SBM.

Santa Barbara: Thome 37534 RSA!; SBBG.

Santa Catalina: Dunkle 1755 AHFH!; Fosberg S4713
LAM!; Thome & Everett 34571 RSA!; SBBG.

San Clemente: Munz 6785 POM!; Raven 18008 RSA!;
SBBG.

Guadalupe: Rempel 759-37 LAM!.

Mesembryanthemum nodiflorum L.

San Miguel: Dunkle 8415 LAM!, AHFH!; SBM.

Santa Rosa: Munz & Hoffmann 1 1724 POM!; SBM.
Santa Cruz: RSA-POM; SBBG.

Anacapa: SBBG; SBM.

San Nicolas: Dunkle 8303 LAM!, AHFH!; Raven &
Thompson 20692 RSA!; Blakley 4131 RSA!; SBM.
Santa Barbara: Dunkle 8147 LAM!; Elmore 310 AHFH!;

Bryan, Dr. & Mrs. s.n. LAM!; Thome 37535 RSA!.
Santa Catalina: Fosberg S4599 LAM!; Dunkle 1812
AHFH!; Thome & Everett 34646 RSA!; SBM.

San Clemente: Dunkle 8121 LAM!; DeBuhr & Wallace
681 RSA!; Raven 17995 RSA!.

Guadalupe: Moran 13757 SD!; Moran 17384.5 (sic) SD!.
Tetragonia tetragonioides (Pall.) Kuntze
San Miguel: Munz & Crow 11814 POM!; SBM.

Santa Rosa: SBM.

Santa Cruz: Balls & Blakley 23756 RSA!.

Amaranthaceae
Amaranthus albus L.

Santa Rosa: Hoffmann s.n. Jun. 12, 1930 SBM!.

Santa Cruz: Dunkle 8591 LAM!, AHFH!; Dunkle 8644
LAM!, AHFH!; Hoffmann s.n. Jun. 14, 1930SBM#5109!;
RSA-POM; SBBG.

Santa Catalina: Fosberg S4509 LAM!; Dunkle 2004
AHFH!; Trask s.n. in Jun. 1896 US!; Trask s.n. in May
1900 NY!.

Amaranthus blitoides Wats.

Santa Rosa: Hoffmann s.n. Jun. 13, 1930 SBM#7735!.
Santa Cruz: Hoffmann s.n. Jul. 1, 1930 SBM# 1845!; RSA-
POM; SBBG.

Amaranthus deflexus L.

Santa Cruz: Raven & Smith 15162 RSA!; Junak 6 1 6 SCIR!;
SBBG.

Anacardiaceae

Lithraea molloides (Kell.) Engler
Santa Catalina: SBBG.

Malosma laurina (Nutt, in T. & G.) Nutt, ex Abrams
Santa Catalina: Trask s.n. in Jun. 1902 LAM!; Fosberg
S5373 LAM!, RSA!; Rusby s.n. Aug. 17, 1915 NY!;
SBM.

San Clemente: Trask 2 1 5 NY; Abrams & Wiggins 393 DS!.
Guadalupe: Rempel 758-37 LAM!.

Rhus integrifolia (Nutt.) Benth. & Hook.

San Miguel: Dunkle 8367 LAM!, AHFH!, RSA!; Voss s.n.

Sep. 2, 1930 POM!; SBBG; SBM.

Santa Rosa: Munz & Crow 1 1643 POM!; Thorne et al.

48853 RSA!; Blakley 3136 RSA!; SBBG; SBM.

Santa Cruz: Fosberg 7625 LAM!, LA!; Dunkle 8595 LAM!,
RSA!; Raven & Smith 15215 RSA!; SBBG; SBM.
Anacapa: Dunkle 7632 AHFH!; Elmore 224 AHFH!;
SBBG; SBM.

Santa Catalina: Fosberg S4286 LAM!; Elmore 439 AHFH!;

Thome 39380 RSA!; SBBG; SBM.

San Clemente: Dunkle 7239 LAM!; Moran 582 LAM!;

DeBuhr & Wallace 721 LAM!; RSA-POM; SBBG.
Guadalupe: Moran 2932 DS!.

Rhus ovata Wats.

Santa Cruz: Fosberg 7686 LAM!, LA!; Thome & Everett
36782 RSA!; Wolf 2782 RSA!; SBBG; SBM.

Santa Catalina: Trask s.n. in Mar. 1902 LAM!; Dunkle
1723 AHFH!, POM!; Raven 17803 RSA!; SBBG; SBM.
Schinus molle L.

Santa Cruz: Blakley 3337 RSA!; SBBG.

Santa Catalina: Thome 36708 RSA!; SBBG.
Toxicodendron radicans (L.) Kuntze ssp. diversilobum (T. &
G.) Thome

San Miguel: Dunkle 8417 LAM!, AHFH!, RSA!; SBBG.
Santa Rosa: Thome et al. 48775 RSA!; Raven, Blakley &
Omduff 14977 RSA!; SBBG; SBM.

Santa Cruz: Fosberg 7709 LAM!, LA!; Elmore 265 AHFH!;
RSA-POM; SBM.

Anacapa: Dunkle 7636 LAM!, AHFH!; SBBG.

Santa Catalina: Fosberg S4301 LAM!; Philbrick & Thome
B67-207 RSA!; SBBG.

San Clemente: Munz 1923 POM!; Raven 17732 RSA!;
SBBG.

Apocynaceae

Asclepias fascicularis Dene, in A. DC.

Santa Cruz: Williams 27 POM!; Balls & Blakley 23712
RSA!; SBBG; SBM.

Santa Catalina: Trask s.n. in Aug. 1902 LAM!; Thome
36678 RSA!; SBBG.

Sarcostemma cynanchoides Dene. ssp. hartwegii (Vail) R.
Holm

Santa Catalina: Trask s.n. in Feb. 1897 US!.

Vinca major L.

Santa Cruz: Thome & Everett 36850 RSA!; SBBG; SBM.
Santa Catalina: Thome & Everett 33465 RSA!.

Araliaceae

Apiastrum angustifolium Nutt, in T. & G.

San Miguel: SBBG.

Santa Rosa: RSA-POM; SBM.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 43

Santa Cruz: Fosberg 7517 LAM!; Fosberg 7612 LAM!;

RSA-POM; SBBG; SBM.

Anacapa: Moran 723 LAM!; SBBG.

San Nicolas: Trask 49 LAM!.

Santa Catalina: Fosberg S4363 LAM!; Dunkle 2073
AHFH!; Fosberg S4717 LAM!; RSA-POM; SBBG.

San Clemente: RSA-POM; SBBG.

Apium graveolens L.

Santa Rosa: Dunkle 8469 LAM!, AHFH!; RSA-POM;
SBBG.

Santa Cruz: SBBG.

San Nicolas: Dunkle 8305 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

San Clemente: DeBuhr & Wallace 718 LAM!.

Berula erecta (Huds.) Cov.

San Miguel: RSA-POM.

Bowlesia incana R. & P.

Santa Rosa: Munz & Crow 11692 LA!; RSA-POM; SBBG;
SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Dunkle 2085 LAM!, AHFH!; Fosberg
S4320 LAM!; Fosberg S7 1 69 LAM!; RSA-POM; SBBG.
San Clemente: Trask 226 NY!; RSA-POM; SBBG.

Caucalis microcarpa H. & A.

Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Fosberg S4339 LAM!, MO!; Fosberg S46 16
LAM!; Detmers s.n. Apr. 13, 1929 USC!; RSA-POM;
SBBG.

San Clemente: RSA-POM.

Conium maculatum L.

Santa Cruz: SBBG; SBM.

San Nicolas: Dunkle 8333 LAM!, AHFH!; Foreman &
Smith 184 LA!; RSA-POM; SBBG.

Daucus pusillus Michx.

San Miguel: Dunkle 8400 LAM!; RSA-POM; SBBG; SBM.
Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Anacapa: SBBG; SBM.

San Nicolas: Dunkle 8310 LAM!; Dunkle 8354 LAM!;

Foreman 230 UC!; RSA-POM; SBBG; SBM.

Santa Barbara: RSA-POM; SBBG.

Santa Catalina: Fosberg S4382 LAM!; Dunkle 1890
AHFH!; Fosberg 4573 LA!, LAM!; RSA-POM; SBBG;
SBM.

San Clemente: Dunkle 7602 LAM!; Dunkle 7300 LAM!;
RSA-POM; SBBG.

Guadalupe: Palmer 34 CM!; Moran 6676 SD!.
Foeniculum vulgare Mill.

Santa Rosa: Dunkle 8460 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

Santa Cruz: Gentry 254 AHFH!; Dunkle 8558 LAM!; RSA-
POM; SBBG; SBM.

San Nicolas: RSA-POM; SBBG.

Santa Catalina: Fosberg S5362 LAM!; RSA-POM; SBBG.
San Clemente: RSA-POM; SBBG.

Lomatium caruifolium (H. & A.) Coult. & Rose
San Miguel: SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Lomatium insulare (Eastw.) Munz

San Nicolas: Trask 51 NY!; Howell 8207 MO!; Foreman
& Smith 174 LA!; CAS; DS; RSA-POM; SBBG; SBM.
San Clemente: Evermann s.n. in 1918 CAS#41543!.
Guadalupe: Moran 6474 CAS!, DS!, LA!, RSA!; Carlquist
473 RSA!; Wiggins & Ernst 138 DS!.

Lomatium utriculatum (Nutt.) Coult. & Rose
Santa Rosa: Munz & Crow 11717 LA!; RSA-POM.

Santa Cruz: Fosberg 7527 LAM!; Fosberg 7658 LAM!;
RSA-POM.

Sanicula arguta Greene ex Coult. & Rose
San Miguel: SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7532 LAM!; Elmore 462 AHFH!;
Ellison s.n. May 12-15, 1929 LA!; RSA-POM; SBBG;
SBM.

Anacapa: SBBG; SBM.

San Nicolas: Trask 94 LAM!; Kanakoff s.n. Apr. 12, 1940
LAM!; Foreman & Smith 197 LA!; RSA-POM; SBBG;
SBM.

Santa Catalina: Templeton 11387 LAM!; Moran 685 LAM!;

Lewis s.n. Mar. 24, 1937 LA!; RSA-POM; SBBG; SBM.
San Clemente: Dunkle 7236 LAM!; Moran 579 LAM!;
RSA-POM; SBBG.

Sanicula crassicaulis Poepp. ex DC.

Santa Cruz: RSA-POM.

Santa Catalina: Trask s.n. in Mar. 1901 NY!; Fosberg S4436
LAM!; Dunkle 2136 LAM!; RSA-POM; SBBG; SBM.
Sanicula hojfmannii (Munz) Bell
San Miguel: RSA-POM.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: Raven & Smith 15174 CAS!; RSA-POM;
SBBG; SBM.

San Nicolas: RSA-POM; SBBG.

Torilis nodosa (L.) Gaertn.

San Miguel: Dunkle 8399 LAM!; RSA-POM; SBBG; SBM.
Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Davidson, A. in May 1914 LAM!; Fosberg
8125 LAM!; Fosberg S4588 SBM!; RSA-POM; SBBG.
Asteraceae

Achillea millefolium L.

San Miguel: Dunkle 8405 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

Santa Rosa: Dunn, N. s.n. May 24, 1939 LA!; RSA-POM;
SBBG; SBM.

Santa Cruz: Clokey 5109 LAM!, LA!; RSA-POM; SBBG;
SBM.

Anacapa: Ellison s.n. May 12-15, 1929 LA!; SBBG; SBM.
San Nicolas: Trask 72 LAM!; Kanakoff s.n. Apr. 14, 1940
LAM!; RSA-POM; SBM.

Santa Barbara: Dunkle 7420 LAM!; Elmore 308 AHFH!;
Bryan, Dr. & Mrs. 3556 LAM!; RSA-POM; SBBG; SBM.

44 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Santa Catalina: Fosberg S4426 LAM!; Fosberg S7168
LAM!; Dunkle 1754 AHFH!; RSA-POM; SBBG; SBM.
San Clemente: Dunkle 7292 LAM!, AHFH!; RSA-POM;
SBBG.

Achyrachaena mollis Schauer

Santa Rosa: Munz & Crow 1 1686 LA!; RSA-POM; SBBG;
SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

San Clemente: Dunkle 7307 AHFH!; RSA-POM; SBBG.
Agoseris apargioides (Less.) Greene ssp. apargioides
Santa Rosa: Dunkle 8499 LAM!, AHFH!; SBM.

Agoseris grandijlora (Nutt.) Greene
San Miguel: SBBG
Santa Rosa: RSA-POM; SBM.

Santa Cruz: Daily 367 SCIR!; SBBG.

Agoseris heterophylla (Nutt.) Greene

Santa Rosa: Hoffmann s.n. Mar. 10, 1932 UC!; RSA-POM;
SBM.

Santa Cruz: RSA-POM.

Guadalupe: Greene s.n. Apr. 24, 1885 GH!.
Amblyopappus pusi/lus H. & A.

San Miguel: Dunkle 8377 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Anacapa: Dunn, N. s.n. May 14, 1932 LA!; RSA-POM;
SBBG.

San Nicolas: Trask s.n. in Apr. 1897 LAM!; Wallace et al.
1621 LAM!; Foreman & Smith 175 LA!; RSA-POM;
SBBG.

Santa Barbara: Dunkle 7434 LAM!; Dunkle 8111 LAM!,
AHFH!, RSA-POM; SBBG; SBM.

Santa Catalina: Trask s.n. in Mar. 1900 LAM!; Fosberg
S4718 LAM!; Dunkle 1886 AHFH!.

San Clemente: DeBuhr & Wallace 693 LAM!; Dunkle 7268
LAM; RSA-POM; SBBG.

Guadalupe: Anthony 258 RSA!; Carlquist 438 RSA!; Mo-
ran 5950 RSA!.

Ambrosia acanthicarpa Hook.

Santa Catalina: RSA-POM.

Ambrosia camphorata (Greene) Payne
Guadalupe: Carlquist 483 RSA!; Moran 6457 RSA!; Greene
s.n. Apr. 24, 1885 CAS#381!.

Ambrosia chamissonis (Less.) Greene

San Miguel: Dunkle 8397 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Dunkle 8599 LAM!; RSA-POM; SBBG; SBM.
Anacapa: Dunkle 7609 LAM!; SBBG; SBM.

San Nicolas: Trask 77 LAM!; Trask 10 CAS!; Raven &
Thompson 20697 LA!; RSA-POM; SBBG; SBM.

Santa Catalina: Dunkle 1896 AHFH!; Fosberg S4879
LAM!; Fosberg S4922 LAM!; RSA-POM; SBBG.

San Clemente: Dunkle 7286 LAM!, AHFH!; RSA-POM;
SBBG.

Ambrosia psilostachya DC. var. californica (Rydb.) Blake in
Tidest.

Santa Catalina: Trask s.n. in Sep. 1899 NY!; Trask s.n. in
Aug. 1902 LAM!; Dunkle 1997 AHFH!; RSA-POM;
SBBG.

Anthemis cotu/a L.

Santa Cruz: Clokey 5113 LAM!; RSA-POM; SBBG; SBM.
Santa Catalina: Dunkle 1978 AHFH!; Trask s.n. in Jun.
1896 US!; RSA-POM; SBBG.

Artemisia californica Less.

San Miguel: Dunkle 8409 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

Santa Rosa: Dunkle 8426 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

Santa Cruz: Yates s.n. in 1 895 LAM!; Dunkle 8538 LAM!,
AHFH!; Clokey 5110 LAM!, LA!; RSA-POM; SBBG;
SBM.

Anacapa: Hilend 281 USC!; Dunkle 7624 LAM!, AHFH!;

Elmore 228 AHFH!; RSA-POM; SBBG; SBM.

Santa Catalina: Dunkle 20 1 5 LAM!; Fosberg 7157 LAM!;

Ewan 10806 LA!; RSA-POM; SBBG; SBM.

San Clemente: DeBuhr & Wallace 713 LAM!; House &
Grumbles s.n. Aug. 5-13, 1930 USC!; Dunkle 7342
LAM!, AHFH!; RSA-POM; SBBG.

Guadalupe: Palmer 48 NY; Moran 6128 SD!.

Artemisia douglasiana Bess, in Hook.

Santa Rosa: Dunkle 8372 LAM!. AHFH!; RSA-POM;
SBBG; SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Trask s.n. in Aug. 1902 LAM!; Fosberg
S5381 LAM!; Dunkle 1998 AHFH!; RSA-POM; SBBG.
Artemisia dracunculus L.

Santa Catalina: Trask s.n. in Aug. 1902 LAM!; Fosberg
S5384 LAM!; SBM.

Artemisia nesiotica Raven

San Nicolas: Trask 71 LAM!, MO!; Trask 71a LAM!, MO!;

Dunkle 8348 LAM!, AHFH!; RSA-POM; SBBG.

Santa Barbara: Bryan, Dr. & Mrs. s.n. Jul. 14, 1922 LAM!;

Dunkle 8126 LAM!; RSA-POM; SBBG; SBM.

San Clemente: Dunkle 7276 LAM!, AHFH!; RSA-POM;
SBBG.

Aster chilensis Nees var. chilensis

Santa Rosa: Hoffmann s.n. Aug. 7, 1930 SBM!.

Santa Cruz: Hoffmann s.n. SBM#10452!; RSA-POM.
Aster exilis Ell.

Santa Cruz: Hoffmann s.n. SBM# 1 04 1 6!; RSA-POM; SBM.
Aster radulinus Gray
Santa Rosa: SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Baccharis douglasii DC.

San Miguel: SBBG.

Santa Rosa: Dunkle 8503 LAM!, AHFH!; Dunkle 8515
LAM!, AHFH!; Epling & Erickson s.n. Aug. 8, 1937 LA!;
RSA-POM; SBBG; SBM.

Santa Cruz: Elmore 281 AHFH!; RSA-POM; SBM.

Santa Catalina: Dunkle 1993 LAM!; Williamsen s.n. Aug.
22, 1 924 USC!; Fosberg S4744 LAM!; RSA-POM; SBBG;
SBM.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 45

Baccharis ernoryi Gray

Santa Catalina: Dunkle 2009 AHFH!; Fosberg 7 1 53 LAM!;
RSA-POM.

Baccharis glutinosa Pers.

Santa Rosa: Dunkle 8504 AHFH!; RSA-POM; SBBG.
Santa Cruz: Moran 758 LAM!; Dunkle 8629 LAM!,
AHFH!; Clokey 5094 LAM!; RSA-POM; SBBG; SBM.
Anacapa: Dunkle 7623 LAM!; SBBG; SBM.

Santa Catalina: Fosberg S4330 LAM!; Fosberg S4797
LAM!; Fosberg S4829 LAM!; RSA-POM; SBBG.

San Clemente: RSA-POM; SBBG.

Baccharis pilularis DC. ssp. consanguinea (DC.) C.B. Wolf
San Miguel: Dunkle 8410 AHFH!; SBBG.

Santa Rosa: Dunkle 8461 LAM!, AHFH!; Epling s.n. in
Nov. 1938 LA!; Dunn, N. s.n. May 24, 1 93 1 LA!; RSA-
POM; SBBG; SBM.

Santa Cruz: Clokey 5093 LAM!; Dunkle 8621 LAM!,
AHFH!; Elmore 285 AHFH!; RSA-POM; SBBG; SBM.
Anacapa: Dunkle 7613 LAM!, AHFH!; SBBG; SBM.

San Nicolas: Trask 79 LAM!; Dunkle 8345 LAM!, AHFH!;

Foreman & Lloyd 133 LA!; RSA-POM; SBBG; SBM.
Santa Barbara: SBBG.

Santa Catalina: Dunkle 202 1 LAM!, AHFH!; Dunkle 2033
LAM!; RSA-POM; SBBG.

San Clemente: RSA-POM.

Baccharis plummerae Gray

Santa Cruz: Dunkle 8579 LAM, AHFH!; Wolf 4 1 27 RSA!;
Thome & Everett 36853 RSA!; SBBG; SBM.
Baeriopsis guadalupensis J. T. Howell
Guadalupe: Rempel 759-37 LAM!; Carlquist 468 RSA!;
Howell 8327A CAS!.

Blennosperma nanum (Hook.) Blake var. nanum
Santa Cruz: SBBG.

Brickelia californica (T. & G.) Gray

Santa Cruz: Yates s.n. in Aug. 1893 LAM!; Dunkle 8543
LAM!; Balls & Blakley 23715 RSA!; SBBG; SBM.
Anacapa: Dunkle 7627 LAM!, AHFH!; Hoffmann s.n. Sep.

22, 1930 POM!; SBBG; SBM.

Santa Catalina: Dunkle 1 729 AHFH!, POM!; Dunkle 2022
LAM!, AHFH!; Thome 35845 RSA!; SBBG.

Centaurea cineraria L.

Santa Catalina: Thome 36239 RSA!; Thome & Everett
33443 RSA!; SBBG.

Centaurea melitensis L.

San Miguel: SBBG.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: Clokey 5166 LAM!, RSA!; Williams 9 POM!;

Blakley 3449 RSA!; SBBG; SBM.

San Nicolas: Raven & Thompson 20769 RSA!; SBBG.
Santa Barbara: Philbrick & McPherson B68-235 RSA!;
Thome 37492 RSA!; SBBG.

Santa Catalina: Dunkle 1845 AHFH!, POM!; Mosquin
3303 US!; Fosberg S4529 LAM!; SBBG.

San Clemente: Raven 17988 RSA!; SBBG.

Guadalupe: Mason 1521 CAS!; Moran 21168 RSA!.
Centaurea solstitialis L.

Santa Cruz: Dunkle 8546 LAM!, AHFH!, NY!; Hoffmann
s.n. Sep. 21, 1930 POM!; Wolf 4 1 69 RSA!; SBBG; SBM.

Santa Catalina: Fosberg 5401 LAM!, NY!, POM!; Thome
36624 RSA!; SBBG.

Chaenactis glabriuscula DC. var. lanulosa (DC.) Hall

Santa Rosa: Munz & Hoffmann 11731 POM!; Blakley 3 1 20
RSA!; Thome et al. 49038 RSA!; SBM.

Chrysanthemum coronarium L.

Santa Cruz: Dunkle 8622 LAM!, AHFH!.

San Nicolas: RSA-POM.

Santa Catalina: Fosberg S4856!.

Chrysanthemum frutescens L.

Santa Catalina: Millspaugh 4728 F!.

Cichorium intybus L.

Santa Cruz: RSA-POM; SBBG; SBM.

Cirsium brevistylum Cronq.

Santa Cruz: Daily 698 SCIR!.

Cirsium californicum Gray

Santa Rosa: Dunkle 8478 LAM!, AHFH!.

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 SBM# 11204!.

Santa Catalina: Davidson s.n. in Jun. 1891 LAM!; Trask
s.n. in May 1 899 US#340043!; Thome & Everett 35015
RSA!; SBM.

Cirsium occidental (Nutt.) Jeps.

San Miguel: Hoffmann s.n. Apr. 10, 1930 LAM!; Hoff-
mann s.n. Jun. 11, 1930 SBM#2758!; Dunkle 8401 LAM!,
AHFH!; RSA-POM; SBBG.

Santa Rosa: Hoffmann s.n. Mar. 10, 1932 SBM#12180!;
Hoffmann s.n. Jul. 9, 1930 SBM# 105 19!; Hoffmann s.n.
May 5, 1932 SBM!; LA; RSA-POM; SBBG.

Santa Cruz: Hoffmann s.n. Mar. 23,1 929 SBM!; Hoffmann
s.n. Jul. 1, 1930 SBM# 1603!; RSA-POM; SBBG.

San Nicolas: RSA-POM; SBBG.

Santa Catalina: Fosberg S4440 LAM!; Fosberg S4876
LAM!, SBM; Dunkle 2470 AHFH!; RSA-POM; SBBG.

San Clemente: Dunkle 7227 LAM!, AHFH!; RSA-POM;
SBBG.

Cirsium ochrocentrum Gray

Santa Catalina: Eastwood 6510 CAS!, US!.

Cirsium proteanum J.T. Howell

Santa Rosa: Hoffmann s.n. May 10, 1932 SBM# 12496!.

Cirsium vulgare (Savi) Ten.

Santa Catalina: Thome 36627 RSA!; SBBG.

Conyza bonariensis (L.) Cronq.

Santa Cruz: Daily 545 SCIR!; SBM.

San Nicolas: Raven & Thompson 20735 RSA!.

Santa Catalina: Thome & Propst 37683 RSA!; Thome
36187 RSA!; SBBG.

San Clemente: Blakley 3653 RSA!; Fosberg & Rainey 55229
RSA!; SBBG.

Conyza canadensis (L.) Cronq.

San Miguel: SBBG.

Santa Rosa: Munz 12810 POM!; Voss s.n. Sep. 4, 1930
POM!; Raven 14978 RSA!; SBM.

Santa Cruz: Hoffmann s.n. Sep. 10, 1931 LAM!; SBM.

San Nicolas: Foreman, Rainey & Evans 77 RSA!; SBBG.

Santa Catalina: Dunkle 2002 AHFH!, POM!; Nuttall 990
POM!; Thome 36663 RSA!.

San Clemente: Raven 17997 RSA!.

46 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Conyza coulteri Gray
Santa Cruz: Dunkle 8579 NY!; SBBG.

Santa Catalina: Millspaugh 4602 F!; McClatchie s.n. Sep.

8, 1893 NY!; Thome & Propst 37679 RSA!.

San Clemente: Moran 7156 RSA!; SBBG.

Coreopsis gigantea (Kell.) Hall
San Miguel: RSA-POM; SBBG; SBM.

Santa Rosa: Dunn, N. s.n. May 15, 1932 LA!; RSA-POM;
SBBG; SBM.

Santa Cruz: Fosberg 7568 LAM!; RSA-POM; SBBG; SBM.
Anacapa: Ellison s.n. May 12-15, 1929 LA!; Dunkle 7607
LAM!, AHFH!; Moran 743 LAM!; SBBG; SBM.

San Nicolas: Trask s.n. in Apr. 1901 LAM!; Dunkle 8360
LAM!, AHFH!; Kanakoffs.n. Apr. 4, 1940 LAM!; RSA-
POM; SBBG.

Santa Barbara: Trask in Apr. 1901 LAM!; Dunkle 7444
LAM!, AHFH!; Elmore 297 AHFH!; RSA-POM; SBBG;
SBM.

Santa Catalina: Fosberg 7161 LAM!; Wolf 363 1 LA!; Mo-
ran 628 LAM!; RSA-POM; SBBG.

Guadalupe: Moran 2923 DS!; Palmer 41 CM!.
Corethrogyne filaginifolia (H. & A.) Nutt. ssp. filaginifolia
San Miguel: SBM.

Santa Rosa: Dunkle 8458 LAM!, AHFH!; RSA-POM; SBM.
Santa Cruz: RSA-POM; SBM.

Anacapa: Dunkle 7633 LAM!, AHFH!; Elmore 220
AHFH!; RSA-POM; SBM.

San Nicolas: Foreman 162 UC!.

Santa Catalina: Trask s.n. in Oct. 1901 LAM!; Dunkle
2452 AHFH!; Fosberg S5375 LAM!; RSA-POM; SBM.
Cotula australis (Sieber ex Spreng.) Hook. f.

Santa Rosa: RSA-POM; SBBG.

Santa Cruz: Fosberg 7579 LAM!; RSA-POM; SBBG; SBM.
Santa Catalina: Fosberg S4506 LAM!; RSA-POM; SBBG.
Cotula coronopifolia L.

San Miguel: SBBG.

Santa Rosa: Elmore 198 AHFH!; Dunkle 8436 LAM!,
AHFH!; Dunn, N. s.n. May 15, 1932 LA!; RSA-POM;
SBM.

Santa Cruz: Clokey 5114 LAM!; Dunkle 8585 LAM!; RSA-
POM; SBBG; SBM.

San Nicolas: Foreman, Evans & Rainey 68 LA!; RSA-
POM; SBBG; SBM.

Santa Catalina: Trask s.n. Mar. 1901 LAM!; Dunkle 1914
AHFH!; Fosberg S4755 LAM!; RSA-POM; SBBG.
Cynara scolymus L.

Santa Cruz: SBM.

Encelia californica Nutt.

Santa Cruz: Fosberg 7531 LAM!, LA!; RSA-POM; SBBG;
SBM.

Anacapa: Dunkle 7672 LAM!; SBBG; SBM.

Santa Catalina: Fosberg S4837 LAM!; Fosberg S4290
LAM!; Lewis s.n. May 24, 1937 LA!; SBBG, SBM.

San Clemente: Dunkle 7280 LAM!; Elmore s.n. Nov. 26,
1929 AHFH!; RSA-POM; SBBG.

Erigeron foliosus Nutt, var .foliosus
San Miguel: SBBG.

Santa Rosa: Elmore 177 AHFH!; RSA-POM; SBM.

Santa Cruz: Clokey 5089 LAM!; Hoffmann s.n. Jun. 15,

1930 LAM!; RSA-POM; SBM.

Anacapa: Moran 727 LAM!.

Santa Catalina: Fosberg S5361 LAM!; Dunkle 1982
AHFH!; RSA-POM; SBBG.

Erigeron foliosus Nutt. var. stenophyllus (Nutt.) Gray
San Miguel: SBBG.

Santa Cruz: RSA-POM; SBBG; SBM.

Anacapa: SBBG.

Erigeron glaucus Ker

San Miguel: Dunkle 8375 LAM!; Elmore 3 1 5 AHFH!; El-
more 332 AHFH!; RSA-POM; SBBG.

Santa Rosa: Dunkle 8480 LAM!; Dunn, N. s.n. May 15,
1932 LA!; RSA-POM; SBBG; SBM.

Santa Cruz: Dunkle 8580 LAM!; Yates s.n. in Aug. 1893
LAM!; Clokey 5091 LAM!, US!; RSA-POM; SBBG;
SBM.

Anacapa: Dunkle 7605 LAM!; Moran 744 LAM!; Dunn,
N. s.n. May 14, 1932 LA!; SBBG; SBM.

Erigeron sanctarum Wats.

Santa Rosa: SBM.

Eriophyllum confertiflorum (DC.) Gray var. confertiflorum
San Miguel: Dunkle 8420 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

Santa Rosa: Dunkle 8483 LAM!; Dunn, N. s.n. May 24,

1931 LA!; Dunn, N. s.n. May 15, 1932 LA!; RSA-POM;
SBBG; SBM.

Santa Cruz: RSA-POM; SBM.

Anacapa: Dunkle 7637 LAM!; Dunn, N. May 14, 1932
LA!; SBBG; SBM.

Santa Catalina: Fosberg 8104 LA!; Fosberg S4397 LAM!;

Dunkle 2163 AHFH!; RSA-POM; SBBG; SBM.

San Clemente: Dunkle 7345 AHFH!; RSA-POM.
Eriophyllum lanatum (Pursh) Forbes var. grandifiorum (Gray)
Jeps.

Guadalupe: Palmer s.n. in 1875 GH!.

Eriophyllum nevinii Gray

Santa Barbara: Dunkle 8101 LAM!; Moran 883 AHFH!;

Elmore 300 AHFH!; RSA-POM; SBBG.

Santa Catalina: Davidson, A. s.n. LAM!; Dunkle 1969
AHFH!; Fosberg S5609 LAM!; RSA-POM; SBBG; SBM.
San Clemente: DeBuhr & Wallace 705 LAM!; Moran 597
LAM!; Nevin s.n. in Apr. 1885 DS!; RSA-POM; SBBG.
Eriophyllum staechadifolium Lag. var. artemisiaefolium
(Less.) Macbr.

San Miguel: SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: SBBG; SBM.

Anacapa: Moran 734 LAM!; SBBG; SBM.

Eriophyllum staechadifolium Lag. var. depressum Greene
San Miguel: RSA-POM.

Santa Rosa: RSA-POM.

Santa Cruz: Clokey 5106 LAM!; Fosberg 7685 LAM!, LA!;

Elmore s.n. Apr. 17, 1936 AHFH!; RSA-POM.
Anacapa: Dunkle 7625 LAM!; RSA-POM.

Evax sparsiflora (Gray) Jeps.

Santa Rosa: RSA-POM; SBM.

San Clemente: RSA-POM.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 47

Filago arizonica Gray
Santa Catalina: RSA-POM.

San Clemente: Dunkle 7317 LAM!; RSA-POM; SBBG.
Guadalupe: Palmer 38 CM!; Palmer 37 (in part) NY!.
Filago californica Nutt.

San Miguel: SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7642 LAM!; Hoffmann s.n. Apr. 1 1,
1931 LAM!; Elmore 443 AHFH!; RSA-POM; SBBG;
SBM.

Anacapa: SBBG.

Santa Catalina: Dunkle 1840 AHFH!; Fosberg S4670
LAM!; Fosberg S4366 LAM!; RSA-POM; SBBG.

San Clemente: Dunkle 7312 LAM!; RSA-POM; SBBG.
Guadalupe: Palmer 37 CM!, MO!, NY!; Palmer 895 ND-
G!; Moran 5662 RSA!.

Filago gallica L.

Santa Cruz; Daily 375 SCIR!; SBBG.

Santa Catalina: RSA-POM.

Gnaphalium beneolens A. Davids.

San Miguel: SBBG.

Santa Rosa: Elmore 21 5 AHFH!; RSA-POM; SBBG; SBM.
Santa Cruz: Clokey 5096 LAM!, US!.

San Nicolas: SBBG.

Santa Catalina: Dunkle 1975 AHFH!.

San Clemente: SBBG.

Gnaphalium bicolor Bioletti

San Miguel: (Hall 1907: Beck 1903 no herbarium cited).
Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Dunkle 86 1 2 LAM!; Elmore s.n. Apr. 18, 1936
USC!; Hoffmann s.n. Apr. 12, 1931 LAM!; RSA-POM;
SBBG; SBM.

Anacapa: SBBG; SBM.

San Nicolas: Dunkle 8358 LAM!, AHFH!; Dunkle 8312
AHFH!; SBBG; SBM.

Santa Catalina: Trask s.n. in May 1896 LAM!; DeBuhr
2583 LAM!; Wolf 3429 US!; RSA-POM; SBBG; SBM.
San Clemente: Moran 595 LAM!, NY!; Trask in Oct. 1902
US!; Elmore 407 AHFH!; RSA-POM; SBBG.
Guadalupe: Moran 13793 RSA!; Moran 18153 RSA!.
Gnaphalium californicum DC.

Santa Rosa: SBM.

Santa Cruz: Clokey 5223 US!; Ellison s.n. May 12-15,
1929 LA!; SBBG; SBM.

Anacapa: SBBG; SBM.

Santa Catalina: Fosberg S4449 LAM!, US!; Dunkle 1974
AHFH!; Blake 9853 US!; RSA-POM; SBBG.
Gnaphalium chilense Spreng.

San Miguel: Dunkle 8381 LAM!, AHFH!; SBBG; SBM.
Santa Rosa: Dunkle 8473 AHFH!; RSA-POM; SBBG;
SBM.

Santa Cruz: Clokey 5097 LAM!, US!; Dunkle 8588 LAM!;

RSA-POM; SBBG; SBM.

Anacapa: SBBG; SBM.

San Nicolas: Dunkle 8312 LAM!; Kanakoff s.n. Apr. 12,
1 940 LAM!; Foreman 1 60 US!; RSA-POM; SBBG; SBM.
Santa Catalina: Fosberg S4338 LAM!, US!; Fosberg S4622
LAM!; Dunkle 1930 AHFH!; RSA-POM; SBBG; SBM.

Guadalupe: Palmer 885 NY!; Moran 6715 RSA!; Moran
17350 RSA!.

Gnaphalium luteo-album L.

San Miguel: SBBG.

Santa Rosa: RSA-POM; SBBG.

Santa Cruz: RSA-POM; SBBG.

San Nicolas: RSA-POM; SBBG.

Santa Catalina: RSA-POM; SBBG.

San Clemente: RSA-POM.

Gnaphalium microcephalum Nutt.

Santa Rosa: Dunkle 8459 LAM!, AHFH!; RSA-POM.
Santa Cruz: RSA-POM; SBBG; SBM.

Anacapa: Dunkle 7620 AHFH!, LAM!.

Santa Catalina: RSA-POM.

Gnaphalium palustre Nutt.

Santa Cruz: Dunkle 8642 LAM!; RSA-POM; SBBG; SBM.
Santa Catalina: Wolf 3598 LAM!, US!; Fosberg S5404
LAM!; Dunkle 1988 AHFH!; RSA-POM; SBBG.

San Clemente: RSA-POM.

Gnaphalium purpureum L.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: Daily 699 SCIR!; SBBG; SBM.

Santa Catalina: RSA-POM; SBBG.

Gnaphalium ramosissimum Nutt.

Santa Cruz: RSA-POM.

Grindelia latifolia Kell. ssp. latifolia
Santa Rosa: Dunkle 8451 LAM!; Moran 783 LAM!, NY!;

Dunn, N. s.n. May 24, 1931 LA!.

Anacapa: Dunkle 7641 LAM!, AHFH!, NY!; Dunkle 7674
LAM!; Kanakoff s.n. Aug. 20, 1940 LAM!; LA.
Grindelia robusta Nutt. var. robusta
Santa Cruz: RSA-POM.

San Nicolas: Wier & Beauchamp s.n. Jun. 29, 1978
UC#1443155!.

Santa Catalina: (Millspaugh & Nuttall 1923: Knopf 493 at
F).

Grindelia stricta DC. ssp. venulosa (Jeps.) Keck
Santa Rosa: SBM.

Haplopappus canus (Gray) Blake
San Clemente: Trask 292 CAS!; SBM.

Guadalupe: Carlquist 455 RSA!; Moran 5971 RSA!.
Haplopappus detonsus (Greene) Raven

Santa Rosa: Dunkle 8493 LAM!; RSA-POM; SBM.

Santa Cruz: Dunkle 8600 LAM!; Brandegee s.n. in 1896
LAM!; Clokey 5087 LAM!, LA!; RSA-POM; SBM.
Anacapa: Dunkle 7614 LAM!; SBM.

Haplopappus palmeri Gray ssp. pachylepis Hall
Santa Catalina: Trask s.n. in Oct. 1900 LAM!.
Haplopappus squarrosus H. & A. ssp. grindelioides (DC.)
Keck

San Miguel: Munz & Crow 1 1805 POM!.

Santa Rosa: Munz & Crow 1 1683 POM!.

Santa Cruz: Hoffmann 217 POM!; Balls & Blakley 23636
RSA!; Wolf 4150 RSA!; SBM; SCIR.

Santa Catalina: Dunkle 2155 AHFH!.

Haplopappus venetus (HBK.) Blake ssp. furfuraceus (Greene)
Hall

San Nicolas: Blakley 4015 SBBG; Foreman 136 UC!.

48 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Santa Catalina: RSA-POM.

San Clemente: RSA-POM.

Haplopappus venetus (HBK.) Blake ssp. sedoides (Greene)
Munz

San Miguel: Dunkle 8376 LAM!; Dunkle 84 1 6 LAM!; RSA-
POM; SBBG.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: Mower s.n. Aug. 21,1 966 LA#9625 1 !; RSA-
POM; SBM.

Anacapa: Dunkle 7643 LAM!; Dunkle 8443 LAM!; SBM.
Haplopappus venetus (HBK.) Blake ssp. vernonioides (Nutt.)
Hall

San Miguel: SBM.

Santa Rosa: Hoffmann s.n. Jun. 13, 1930 SBM#2691!;

Epling & Erickson s.n. Aug. 8, 1937 LA!.

Santa Cruz: Hoffmann s.n. Nov. 9, 1930 SBM#10816!;

Mower s.n. Aug. 22, 1966 LA#96256!; RSA-POM.
Anacapa: Dunkle 7676 AHFH!; Elmore 246 AHFH!; Hoff-
mann s.n. Sep. 22, 1930 SBM#10278!.

San Nicolas: Wallace et al. 1612 LAM!; Wallace et al. 1615
LAM!; Foreman 215 LA!; RSA-POM; SBBG; SBM.
Santa Catalina: Dunkle 2024 LAM!; Fosberg S4855 LAM!;

Fosberg 7151 LAM!; RSA-POM.

San Clemente: RSA-POM.

Helenium puberulum DC.

Santa Catalina: RSA-POM.

Helianthus annuus L. ssp. lenticularis (Dougl.) Ckll.

Santa Cruz: Laughrin 528 SCIR!.

Santa Catalina: Trask s.n. in Mar. 1901 F!; Nuttall 569 F!.
Hemizonia Clementina Bdg.

Anacapa: Dunkle 7642 LAM!; Hoffmann s.n. Jun. 16, 1930
LAM!; Ellison s.n. May 12-15, 1929 LA!; SBBG; SBM.
San Nicolas: Trask 80 LAM!; Foreman 214 US!; Raven
& Thompson 20773 LA!; RSA-POM; SBBG; SBM.
Santa Barbara: Dunkle 8113 LAM!; Bryan, Dr. & Mrs. s.n.
Jul. 1, 1922 LAM!, US!; Blakley 4799 US!; RSA-POM;
SBBG; SBM.

Santa Catalina: Fosberg S4887 LAM!; Nuttall 1 95 F!; East-
wood 6492 US!; RSA-POM; SBBG; SBM.

San Clemente: Dunkle 7277 LAM!; Raven 17836 LA!;
Meams 4046 US!; RSA-POM; SBBG.

Hemizonia fasciculata (DC.) T. & G.

Santa Rosa: Dunkle 8452 LAM!; SBBG; SBM.

Santa Cruz: Dunkle 8582 LAM!; Greene s.n. in Jul. -Aug.

1886 US!; RSA-POM; SBBG; SBM.

Anacapa: Dunkle 7675 LAM!; SBBG.

Santa Barbara: Dunkle 8 1 37 LAM!; Bryan, Dr. & Mrs. s.n.
LAM!; SBBG.

Santa Catalina: Fosberg S4930 LAM!, NY!; Trask s.n. in
May 1898 US!; Dunlavy s.n. May 4, 1934 LA!; RSA-
POM; SBBG.

San Clemente: Trask 195 US!; RSA-POM; SBBG.
Hemizonia fitchii Gray

Santa Cruz: Nixon 1773 SCIR!; RSA-POM; SBBG.
Hemizonia frutescens Gray
Guadalupe: Palmer 42 MO!, NY!, CM!.

Hemizonia greeneana Rose ssp. greeneana

Guadalupe: Rempel 758-37 LAM!; Moran 18160 LAM!;
Palmer 865 NY!; LA; MO; RSA-POM.

Hemizonia increscens (Hall ex Keck) Tanowitz ssp. incres-
cens

Santa Rosa: Brandegee s.n. in Jun. 1888 US!; SBM.

Santa Cruz: SBM.

Hemizonia palmeri Rose

Guadalupe: Moran 17337 MO!; Lindsay 43082 RSA!;
Carlquist 470 RSA!.

Heterotheca grandiflora Nutt.

Santa Cruz: RSA-POM; SBBG.

San Nicolas: RSA-POM.

Santa Catalina: Dunkle 1 976 LAM!; Fosberg S4302 LAM!;

Fosberg 4447 LAM!; RSA-POM; SBBG.

San Clemente: DeBuhr & Wallace 688 LAM!; RSA-POM;
SBBG.

Hieraceum argutum Nutt. ssp. argutum

Santa Rosa: Dunkle 8482 AHFH!; RSA-POM; SBM.
Santa Cruz: Clokey 5088 LAM!; RSA-POM; SBBG; SBM.
Hypochoeris glabra L.

Santa Rosa: Thome et al. 48782 RSA!; Raven, Blakley &
Omduff 14885 RSA!; Munz&Crow 1 1640 POM!; SBBG;
SBM.

Santa Cruz: Sauer & Hobbs 6029 LA!; Pierson 11031 RSA!;

Clokey 5081 POM!; SBBG; SBM.

Santa Catalina: Thome 36305 RSA!; SBBG.

San Clemente: Raven 1 7693 RSA!; SBBG.

Guadalupe: Moran 13787 RSA!; Norris, K. S. s.n. May 1,
1951 LA!.

Jaumea carnosa (Less.) Gray
San Miguel: SBM.

Santa Rosa: Dunkle 8492 LAM!; Munz & Hoffmann 1 1726
POM!; SBM.

Santa Cruz: SBM.

Santa Catalina: Fosberg S4895 LAM!, POM!; Thome 45090
RSA!; SBM!.

Lactuca serriola L.

Santa Rosa: SBM.

Santa Cruz: Daily 404 SCIR!; SBBG.

San Nicolas: SBBG.

Santa Catalina: Thorne 36276 RSA!; SBBG.

Lasthenia californica DC. ex Lindl.

San Miguel: Munz & Voss 1 1873 POM!; SBBG; SBM.
Santa Rosa: Hoffmann s.n. May 10, 1932 POM!; Raven,
Blakley & Omduff 14921 RSA!; Thorne et al. 48859
RSA!; SBBG; SBM.

Santa Cruz: Fosberg 7523 LAM!; Elmore 468 AHFH!;

Raven & Smith 15302 RSA!; SBBG, SBM.

Anacapa: Dunn, N. s.n. May 14, 1932 LA!; SBBG; SBM.
San Nicolas: Dunkle 8351 LAM!; Kanakoff s.n. Apr. 12,
1940 LAM!; SBBG; SBM.

Santa Barbara: Dunkle 7402 LAM!, AHFH!; Blakley 5695
RSA!; Dunkle 7467 AHFH!; SBBG; SBM.

Santa Catalina: Dunkle 1742 AHFH!, POM!; Fosberg
S4376 LAM!; Wolf 3518 POM!; SBBG; SBM.

San Clemente: Dunkle 7261 LAM!; Elmore 400 AHFH!;
Raven 17193 RSA!; CAS; SBBG.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 49

Guadalupe: Thobum, Greene & Wing s.n. in Jul. 1897
DS!; Wiggins & Ernst 119 DS!; Greene s.n. in 1885
DS#205127!; RSA-POM.

Lasthenia coronaria (Nutt.) Ornduff
Guadalupe: Anthony s.n. in May-Jun. 1897 UC!.
Lasthenia glabrata Lindl. ssp. coulteri (Gray) Ornduff
Santa Rosa: Hoffmann s.n. Apr. 8, 1930 POM!; Raven
14990 RSA!

Layia glandulosa (Hook.) H. & A. ssp. glandulosa
Santa Cruz: RSA-POM.

Layia platyglossa (F. & M.) Gray ssp. campestris Keck
San Miguel: Youngberg s.n. May 28, 1939 POM!; Munz
& Crow 11824 POM!; SBBG; SBM.

Santa Rosa: Dunn, N. May 24, 1931 LA!; Thome et al.

48858 RSA!; Munz & Crow 1 1 703 POM!; SBBG; SBM.
Santa Cruz: Fosberg 7602 LAM!; Moran 752 LAM!, RSA!;

Clokey 5103 NY!; LA; SBBG; SBM.

Santa Catalina: Fosberg S4814 LAM!, NY!; Dunkle 1834
AHFH!; Thome 35928 RSA!; SBBG; SBM.

San Clemente: Dunkle 7233 LAM!; Thome 42948 RSA!,
NY!; Raven 17331 RSA!; SBBG.

Guadalupe: Anthony 251 LIS!.

Layia platyglossa (F. & M.) Gray ssp. platyglossa

Santa Rosa: Moran 804 LAM!, NY!; Elmore 202 AHFH!;
Dunkle 8428 AHFH!; SBM.

Lepidospartium squamatum (Gray) Gray
Santa Cruz: Balls & Blakley 23750 RSA!; SBBG.

Madia exigua (Sm.) Gray

Santa Cruz: Hoffmann s.n. Apr. 10, 1931 LA!; Clokey 5102
POM!; Raven & Smith 15228 RSA!; SBBG; SBM.
Santa Catalina: Davidson A. s.n. Jun. 26, 1891 LAM!;
Fosberg S4833 LAM!, POM!; Raven 17812 RSA!; SBBG;
SBM.

Madia gracilis (Sm.) Keck ssp. gracilis

Santa Cruz: Clokey 5101 LAM!, POM!; Raven & Smith
15178 RSA!; Thome & Everett 36847 RSA!; SBBG;
SBM.

Santa Catalina: Fosberg S4427 LAM!; Thome 36857 RSA!,
LA!; Raven 17818 RSA!; SBBG; SBM.

Madia sativa Mol.

Santa Cruz: Dunkle 8578 LAM!, AHFH!; SBBG.

Santa Catalina: Dunkle 2450 AHFH!; Trask s.n. US!;

Thome & Everett 35005 RSA!; SBBG.

San Clemente: Raven 17983 RSA!.

Malacothrix cleve/andii Gray

Guadalupe: Palmer 51 NY!, CM!.

Malacothrix coulteri Harv. & Gray var. cognata Jeps.

Santa Rosa: Hoffmann 699 POM!; SBM.

Santa Cruz: Fosberg 7535 LAM!; SBM.

Malacothrix foliosa Gray

San Clemente: Trask 21 3 NY!; Dunkle 721 1 LAM!, AHFH!,
NY!; Nevin & Lyon s.n. in Apr. 1885 CAS#731!,
DS#1 17575!; RSA-POM; SBBG.

Malacothrix incana (Nutt.) T. & G.

San Miguel: Dunkle 8412 LAM!, MO!, NY!; Elmore 327
AHFH!; Yates s.n. LAM!; SBBG; SBM.

Santa Rosa: Thome et al. 48912 RSA!; Brandegee s.n. in
1888 GH!; Dunkle 8462 LAM!.

Santa Cruz: Greene s.n. in Jul.-Aug. 1886 ND-G#065 1 69!,
NY!.

San Nicolas: Wier & Beauchamp s.n. Jul. 4, 1968
RSA#289130!.

San Clemente: (Raven 1963: Murbarger 189 at UC.)
Malacothrix indecora Greene
San Miguel: SBM.

Santa Cruz: Greene s.n. in Jul.-Aug. 1886 NY!, MO!.
Malacothrix “A” (Davis 1980)

Santa Barbara: Dunkle 8105 LAM!; Dunkle 8133 LAM!,
NY!; Elmore 370 AHFH!; RSA-POM!.

Malacothrix “C” (Davis 1980)

San Nicolas: Trask s.n. in Apr. 1 897 MO!; Foreman, Evans
& Rainey 80 UC!; Wier & Beauchamp s.n. Jul. 3, 1978
RSA!.

Malacothrix saxatilis (Nutt.) T. & G. var. implicata (Eastw.)
Hall

San Miguel: Dunkle 8388 LAM!, NY!; Moran 3439 POM!,
NY!; Hoffmann s.n. Apr. 10, 1930 LA!; SBBG; SBM.
Santa Rosa: Dunkle 8433 LAM!; Blakley 3192 RSA!; Dunn,
N. s.n. May 24, 1931 LA!; SBBG; SBM.

Santa Cruz: Clokey 5080 NY!; Raven & Smith 15127 RSA!;

Eastwood 6416 NY!; SBBG; SBM.

Anacapa: Dunkle 7608 LAM!, NY!; Elmore 248 AHFH!;

Ellison s.n. May 12-15, 1929 LA!; SBBG; SBM.

San Nicolas: Trask 65 LAM!, NY!; Trask 64 MO!; Raven
& Thompson 20760 RSA!; SBBG; SBM.

Malacothrix saxatilis (Nutt.) T. & G. var. tenuifolia (Nutt.)
Gray

Santa Catalina: Fosberg S4423 LAM!, NY!; Thome 36469
RSA!; Nuttall 1010 POM!; SBBG; SBM.

Malacothrix similis Davis & Raven

San Miguel: (Davis 1982: Greene s.n. in Sep. 1886
CAS#734)

Santa Cruz: (Davis & Raven 1962: Brandegee s.n. in 1888
at UC.)

Malacothrix squalida Greene

Santa Cruz: Greene s.n. in Jul.-Aug. 1886 CAS#735!; NY!.
Anacapa: SBBG.

Matricaria matricarioides (Less.) Porter

Santa Rosa: Blakley 3 1 93 RSA!; Thome et al. 48719 RSA!;
SBBG; SBM.

Santa Cruz: Daily 37 SCIR!; SBBG; SBM.

Santa Catalina: Fosberg S4507 LAM!, POM!; Dunkle 245 1
AHFH!; SBBG.

Guadalupe: Palmer 47 CM!; Wiggins & Ernst 1 13 SD!.
Micropus californicus F. & M.

Santa Rosa: Sweet s.n. Apr. 15, 1935 POM!; Thome et al.
48997B RSA!; SBM.

Santa Cruz: Raven & Smith 15189 RSA!; Blakley 3295
RSA!; Munz & Crow 11515 POM!; SBBG; SBM.
Microseris douglasii (DC.) Sch.-Bip. ssp. douglasii
San Nicolas: Raven & Thompson 20767 RSA!.

San Clemente: Thome 42789 RSA!.

Microseris douglasii (DC.) Sch.-Bip. ssp. platycarpha (Gray)
Chamb.

Santa Catalina: Thome 35778 RSA!; Thome 35935a RSA!;
SBBG.

50 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San Clemente: Munz 6739 POM!; Dunkle 7364 AHFH!;
Raven 17199 RSA!; SBBG.

Microseris douglasii (DC.) Sch.-Bip. ssp. tenella (Gray)
Chamb.

San Miguel: SBM.

Santa Rosa: SBM.

Santa Cruz: SBM.

Microseris elegans Greene ex Gray
San Miguel: SBM.

Santa Cruz: SBBG; SBM.

San Clemente: Thome 42819b RSA!.

Microseris heterocarpa (Nutt.) Chamb.

Santa Rosa: Thome 48980 RSA!; Raven, Blakley & Om-
duff 14908 RSA!; Hoffmann s.n. Mar. 8, 1932 UC!;
SBBG.

Santa Cruz: Brandegee s.n. in Apr. 1888 UC!.

Santa Catalina: Brandegee, K. s.n. UC!; Thome 36347
RSA!, UC!; SBBG.

San Clemente: Trask 235 NY!; Dunkle 7325 AHFH!; Ra-
ven 17206 RSA!; SBBG.

Guadalupe: Brandegee s.n. May 20, 1897 UC!; Wiggins &
Ernst 206 UC!; Moran 17362 RSA!.

Microseris linearifolia (DC.) Sch.-Bip.

Santa Rosa: Blakley & Smith 3068 RSA!; Hoffmann s.n.

in Apr. 1930 CAS#178617!; SBBG; SBM.

Santa Cruz: Blakley 3356 RSA!; Hoffmann s.n. Apr. 1 1,
1931 LA!; Abrams & Wiggins 50 CAS!; SBBG; SBM.
Anacapa: SBBG; SBM.

San Nicolas: Trask 66 ND-G#063915!, NY!.

Santa Barbara: (Philbrick 1972: Philbrick & McPherson
B68-216)

Santa Catalina: Fosberg S4419 LAM!, NY!; Thome 35805
RSA!; Dunkle 1823 AHFH!; SBBG.

San Clemente: Dunkle 7296 LAM!, AHFH!; Thome 42790
RSA!; Raven 17612 RSA!; SBBG.

Guadalupe: Palmer 50 MO!; Anthony 243 MO!; Moran
17361!.

Pentachaeta lyonii Gray

Santa Catalina: Fosberg S5416 LAM!; RSA-POM.
Perezia microcephala (DC.) Gray
Santa Rosa: Dunkle 8424 LAM!; Dunn 1303 LA!; Munz
& Crow 11612 POM!; SBBG; SBM.

Santa Cruz: Dunkle 8626 LAM!; Elmore 263 AHFH!;

Blakley 3306 RSA!; SBBG; SBM.

Santa Catalina: Fosberg S5380 LAM!; Diehl 258 POM!;
Thome & Everett 34890 RSA!; USC; SBBG; SBM.
Perityle emoryi Torr. in Emory

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Dunkle 86 1 7 LAM!, NY!; Greene s.n. in 1 886
NY; Williams 52 POM!; SBBG; SBM.

Anacapa: SBBG; SBM.

Santa Barbara: Dunkle 8104 LAM!, NY!; Thorne 37549
RSA!; Blakley 5626 RSA!; SBBG.

Santa Catalina: Thome 35754 RSA!; Trask s.n. in Mar.

1899 NY; Dunkle 1954 LAM!, AHFH!; SBBG; SBM.
San Clemente: Dunkle 7335 LAM!; Trask 255 NY!; Raven
17323 RSA!; SBBG.

Guadalupe: Palmer 44 NY!, CM!; Carlquist 439 RSA!;
Moran 5628 RSA!.

Perityle incana Gray

Guadalupe: Rempel 758-37 LAM!; Rempel 759-37 LAM!;
Palmer 43 CM!; RSA-POM; SBM.

Picris echioides L.

Santa Catalina: Thome 36450 RSA!; Thorne 36890 RSA!;
Thorne 36664 RSA!; SBBG.

Pluchea odorata (L.) Cass.

Santa Cruz: Hoffmann & Cooke s.n. Sep. 20, 1930 POM!;
SBBG; SBM.

Santa Catalina: Thorne 36677 RSA!; Dunkle 1 993 AHFH!;
SBBG.

Pluchea sericea (Nutt.) Cov.

Santa Catalina: Fosberg S4479 LAM!, POM!.
Psilocarphus tenellus Nutt. var. tenellus
Santa Rosa: Sweet s.n. Apr. 15, 1935 POM!; Raven 14953
RSA!; Hoffmann s.n. Jun. 1 5, 1 930 LAM!; SBBG; SBM.
Santa Cruz: Blakley 3342 RSA!; Thome & Everett 36779
RSA!; Raven & Smith 15211 RSA!; SBBG; SBM.

Santa Catalina: Thorne 35987 RSA!.

San Clemente: Thome 36087 RSA!; Thome 42974 RSA!;
Raven 17664 RSA!.

Rafnesquia californica Nutt.

Santa Rosa: SBM.

Santa Cruz: Brandegee s.n. in 1888 RSA!; SBBG; SBM.
Anacapa: SBBG.

Santa Barbara: Thome 3751 1 RSA!; SBBG.

Santa Catalina: Fosberg S4537 LAM!; Dunkle 2149
AHFH!; Thome & Everett 34888 RSA!; SBBG.

San Clemente: Raven 17608 RSA!; Raven 17244 RSA!;
Thome 42852 RSA!; SBBG.

Senecio aphanactis Greene
Santa Rosa: SBM.

Santa Cruz: Pierson s.n. Mar. 25, 1934 POM!; Wolf 2780
RSA!; Wolf 2805 RSA!; SBM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!.

Senecio douglasii DC. var. douglasii
Santa Cruz: Balls & Blakley 23665 RSA!; Balls & Blakley
23633 RSA!; Hoffmann s.n. Jul. 20, 1930 POM!; SBBG;
SBM.

Santa Catalina: Nuttall 794 F!.

San Clemente: Dunkle 7227 LAM!.

Senecio lyonii Gray

Santa Catalina: Grant & Wheeler 6134 LAM!, POM!;

Thome 39388 RSA!; Raven 17817 RSA!; SBBG; SBM.
San Clemente: Dunkle 7349 LAM!; Moran 589 LAM!;
Raven 17161 RSA!.

Senecio palmeri Gray

Guadalupe: Palmer 49 CM!; Franceschi 10 SBM!.

Senecio vulgaris L.

San Miguel: SBBG.

Santa Cruz: Raven & Smith 15222 RSA!; SBBG; SBM.
Anacapa: SBBG.

San Nicolas: Thome et al. 52350 RSA!.

Santa Catalina: Thome 36209 RSA!; Thome, Rollins,
Propst & Carolin 36751 RSA!; SBBG.

San Clemente: Raven 17230 RSA!; SBBG.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 51

Silybum marianum (L.) Gaertn.

Santa Rosa: Thome et al. 48803 RSA!.

Santa Cruz: Dunkle 8539 LAM!, AHFH!; Elmore 264
AHFH!; Clokey 5115 LAM!, POM!; SBBG; SBM.
Santa Barbara: Philbrick & Ricker B69-48 RSA!.

Santa Catalina: Fosberg S4931 LAM!; Thome & Everett
35047 RSA!; SBBG.

Solidago californica Nutt.

Santa Rosa: Dunkle 8444 LAM!; Epling & Erickson s.n.

Aug. 8, 1937 LA!; Voss s.n. Sep. 4, 1930 POM!; SBM.
Santa Cruz: Hoffmann s.n. Sep. 9, 1931 LAM!; Hoffmann
s.n. Sep. 20, 1930 POM!; Blakley 23690 RSA!; SBBG;
SBM.

Santa Catalina: Dunkle 2001 AHFH!, POM!; Thome 36703
RSA!; SBBG.

Sonchus asper (L.) Hill

San Miguel: Dunkle 8396 LAM!; Hoffmann s.n. Mar. 25,
1927 SBM#1069!; SBBG.

Santa Rosa: Dunkle 8521 AHFH!; RSA-POM; SBBG;
SBM.

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 LAM!; Clokey
5079 NY!; RSA-POM; SBBG; SBM.

Anacapa: SBBG.

San Nicolas: Trask s.n. in Apr. 1901 LAM!; Dunkle 8330
AHFH!; Kanakoff s.n. Apr. 12, 1940 LAM!.

Santa Barbara: Bryan, Dr. & Mrs. s.n. LAM!; SBM.

Santa Catalina: Fosberg S4340 LAM!; Fosberg S4620
LAM!; Dunkle 2095 AHFH!; RSA-POM; SBBG.

San Clemente: Dunkle 7209 LAM!, AHFH!; RSA-POM;
SBBG.

Sonchus oleraceus L.

San Miguel: Dunkle 8421 LAM!; SBBG; SBM.

Santa Rosa: SBBG; SBM.

Santa Cruz: Dunkle 8589 LAM!; RSA-POM; SBBG; SBM.
Anacapa: Dunkle 7649 AHFH!; SBBG; SBM.

San Nicolas; Dunkle 8306 LAM!, AHFH!, NY!; Foreman,
Evans & Rainey 25 LA!; RSA-POM; SBBG.

Santa Barbara: Elmore 304 AHFH!; Dunkle 8131 LAM!,
NY!; Dunkle 7450 LAM!, AHFH!, NY!; RSA-POM;
SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 NY!; Dunkle 2468
AHFH!; Knopf 224 F!; RSA-POM; SBBG.

San Clemente: Trask 353 NY!; RSA-POM; SBBG.
Guadalupe: Palmer 51 NY!; Palmer 871 NY!, LIS!; Moran
6615 RSA!.

Sonchus tenerrimus L.

San Nicolas: Trask s.n. in Apr. 1900 LAM!; Trask 23 US!;
Trask 67 MO!.

Santa Barbara: (Philbrick 1972: Philbrick & McPherson
B68-247)

Santa Catalina: Trask s.n. in Mar. 1901 NY!; Millspaugh
4544 F!.

San Clemente: Trask 355 NY!, US!; Trask 356 NY!, US!.
Guadalupe; Palmer 873 NY!, US!; Moran 17416 SD!;
Moran 13763 SD!.

Stephanomeria blairii M. & J.

San Clemente: Raven 17310 RSA!; Dunkle 7353 LAM!,
AHFH!; House & Grumbles s.n. USC!; SBBG.

Stephanomeria cichoriacea Gray
Santa Rosa: SBM.

Santa Cruz: Hoffmann s.n. Jun. 15, 1930 LAM!; Dunkle
8531 LAM!; Fosberg 7561 LAM!; RSA-POM; SBBG;
SBM.

Stephanomeria diegensis Gottlieb
Santa Rosa: Epling & Erickson s.n. Aug. 8, 1937 LA!.
Santa Catalina: Piehl 62515 SBBG!; Millspaugh 4534 F!;
Fosberg S7 147 LAM!

San Clemente: Piehl 631058 SBBG!; Dunkle 7354 LAM!,
AHFH!, SBBG!; Raven 18015 SBBG; Abrams & Wig-
gins 344 CAS!; RSA-POM.

Stephanomeria exigua Nutt. ssp. coronaria (Greene) Gott-
lieb

San Miguel: Greene s.n. in 1886 ND-G#001750!.

Santa Cruz: RSA-POM.

Stephanomeria guadalupensis Bdg.

Guadalupe: Rempel s.n. Jul. 19, 1937 LAM!; Carlquist
457 RSA!; Moran 15119 RSA!.

Stephanomeria virgata Benth. ssp. virgata
Santa Rosa: Dunkle 8477 LAM!; RSA-POM; SBBG; SBM.
Santa Cruz: Dunkle 8625 LAM!; RSA-POM; SBBG; SBM.
Santa Catalina: RSA-POM; SBBG.

San Clemente: RSA-POM; SBBG.

Styloc/ine gnaphalioides Nutt.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg S7684 LAM!; RSA-POM; SBBG;
SBM.

Santa Catalina; Grant 1256 LAM!; Fosberg S4601 LAM!;
Dunkle 2145 AHFH!; RSA-POM; SBBG! SBM.
Taraxacum laevigatum (Willd.) DC.

Santa Cruz: RSA-POM.

Santa Catalina: RSA-POM.

Taraxacum officinale Wiggers
Santa Cruz: Daily 610 SCIR!; SBBG.

Santa Catalina: SBBG.

Thelesperma megapotamicum (Spreng.) Kuntze
Santa Catalina: Nuttall 602 F!.

Tragopogon porrifolius L.

Santa Cruz: SBBG; SBM.

Venegasia carpesioides DC.

Santa Rosa: Moran 814 LAM!; RSA-POM; SBBG; SBM.
Santa Cruz: Dunkle 8608 LAM!; Fosberg 7697 LAM!, LA!;
Clokey 5108 LAM!; RSA-POM; SBBG; SBM.
Xanthium spinosum L.

San Miguel: Dunkle 8364 LAM!; RSA-POM; SBBG; SBM.
Santa Cruz: Daily 148 SCIR!; SBBG.

Santa Barbara: Abrams & Wiggins 39 DS!.

Santa Catalina: Trask s.n. in Aug. 1902 LAM!, NY!; Trask
s.n. in Aug. 1 897 US!; Fosberg S4547 LAM!; CAS; RSA-
POM; SBBG.

Xanthium strumarium L. var. canadense (Mill.) T. & G.
Santa Catalina: Dunkle 2437 AHFH!; Fosberg S4897
LAM!; RSA-POM.

Bataceae

Bat is maritima L.

San Clemente: Nevin s.n. in 1885 DS#81927!.

52 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Berberidaceae

Berberis pinnata Lag. ssp. insularis Munz
Santa Rosa: RSA-POM; SBM.

Santa Cruz: Hoffmann 1 89 LAM!, LA!; RSA-POM; SBBG;
SBM.

Anacapa: Dunkle 7668 LAM!; RSA-POM; SBBG.
Boraginaceae

Amsinckia intermedia F. & M.

San Miguel: Schuyler 21 LAM!; Munz & Norris 11790
POM!, GH!; SBBG; SBM.

Santa Rosa: Moran 779 LAM!; Elmore 2 1 2 AHFH!; Thome
et al. 48732 RSA!; SBBG; SBM.

Santa Cruz: Fosberg 7556 LAM!; Fosberg 7520 LAM!;

Elmore s.n. Apr. 18, 1936 LAM!; SBBG; SBM.
Anacapa: SBBG; SBM.

Santa Barbara: Dunkle 7461 LAM!, DS!, NY!; Dunkle
8112 LAM!; Thome 37537 RSA!; SBBG; SBM.

Santa Catalina: Trask s.n. in Apr. 1898 US!; Millspaugh
4700 F!; Templeton 11391 LAM!; CAS; DS; RSA-POM;
SBBG; SBM.

San Clemente: Moran 569 LAM!, DS!, GH!, NY!, RSA!;
Elmore 427 AHFH!; Trask 248 US!; CAS; SBBG.
Amsinckia menziesii (Lehm.) Nels. & Macbr.

Santa Cruz: Moran 773 LAM!; Raven & Smith 15261
RSA!; SBBG.

Santa Catalina: Dunkle 2060 AHFH!; Wolf 3622 RSA!;
SBBG.

Guadalupe: Palmer 69 GH!, MO!, NY!; Moran 668 1 RSA!;
Howell 8329 CAS!.

Amsinckia spectabilis F. & M. var. nicolai (Jeps.) Jtn. ex
Munz

San Miguel: Munz & Crow 1 1789 POM!; SBM.

San Nicolas: Kanakoff s.n. Apr. 22, 1940 LAM!; Trask 58
LAM!; Dunkle 8339 LAM!, RSA!; CAS; US.

San Clemente: Munz 6771 POM!; Thome 42942 RSA!.
Amsinckia spectabilis F. & M. var. spectabilis

San Miguel: Greene s.n. in Sep. 1886 ND-G#Q42845!;
SBBG; SBM.

Santa Rosa: Thome et al. 49040 RSA!; Munz & Crow
1 1570 POM!, US!; Raven 14986 RSA!; SBBG; SBM.
Santa Cruz: Webster, Axelrod et al. 84 RSA!; Raven &
Smith 15309 RSA!; SBBG; SBM.

Anacapa: SBBG; SBM.

San Nicolas: Kanakoff s.n. Apr. 16, 1940 LAM!; Trask s.n.

in Apr. 1897 US!; Trask 59 GH!; CAS.

Santa Barbara: Dunkle 7408 AHFH!.

San Clemente: Raven 17262 RSA!; SBBG.

Cryptantha clevelandii Greene var. c/evelandii
San Miguel: Schuyler s.n. Apr. 1, 1976 LAM!; Elmore 312
AHFH!; Munz & Crow 1 1823 POM!; SBBG; SBM.
Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Clokey 5037 NY!; Clokey 5038 NY!; RSA-
POM; SBBG; SBM.

Anacapa: SBBG; SBM.

San Nicolas: RSA-POM.

Santa Barbara: Dunkle 7446 LAM!, AHFH!; Dunkle 7428
LAM!; RSA-POM; SBBG; SBM.

Santa Catalina: Fosberg 1 0647 LAM!, LA!; Trask s.n. Mar.

1897 NY!; Millspaugh 4606 F!; RSA-POM; SBBG; SBM.
San Clemente: RSA-POM.

Cryptantha clevelandii Greene var. jlorosa Jtn.

San Miguel: Hoffmann s.n. Apr. 10, 1930 LAM!; Young-
berg, F. s.n. May 28, 1938 POM#259646!.

Santa Rosa: Hoffmann s.n. Apr. 18, 1929 SBM#5996!;

Munz & Crow 11710 POM!.

Santa Cruz: Fosberg 7619 LAM!, LA!; Hoffmann s.n. Jun.

15, 1930 LAM!; Abrams & Wiggins 110 NY!; POM.
Anacapa: Hoffmann s.n. Mar. 16, 1929 SBM!.

San Nicolas: Kanakoff s.n. Apr. 12, 1940 LAM!.

Santa Barbara: Dunkle 7447 LAM!; Dunkle 7428 LAM!.
Santa Catalina: Dunkle 2097 LAM!; Fosberg S4750 LAM!;
Hasse 4156 NY!.

Cryptantha foliosa (Greene) Greene
Guadalupe: Greene s.n. Apr. 26, 1885 NY!; Palmer 68
CM!, MO!; Moran 5954 LA!, NY!, RSA!.

Cryptantha intermedia (Gray) Greene

Santa Catalina: Trask s.n. May 1 895 MO#25 1 7956!; Trask
s.n. in Mar. 1901 US!; Millspaugh 4744 F!; RSA-POM;
SBBG.

San Clemente: Dunkle 7260 LAM!, AHFH!; Trask 176
NY!; Trask 277 NY!; RSA-POM; SBBG.

Cryptantha leiocarpa (F. & M.) Greene
Santa Rosa: Raven 14998 UC!.

Cryptantha maritima (Greene) Greene
San Nicolas: Trask 57 LAM!, GH!, NY!; Trask 56 GH!;

Dunkle 8353 AHFH!; SBBG.

Santa Barbara: Dunkle 7403 LAM!, NY!; RSA!; Dunkle
8106 AHFH!; Philbrick & Benedict B66-362 RSA!;
SBBG.

Santa Catalina: Trask s.n. in Mar. 1900 NY!; Knopf 143
F!; Thome 35855 RSA!; SBBG.

San Clemente: Trask 221 NY!; Munz 6746 POM!; Raven
17318 RSA!; SBBG.

Guadalupe: Palmer 67 CM!, MO!, NY!; Greene s.n. Apr.
26, 1885 NY; Carlquist 449 RSA!; CAS; DS.
Cryptantha micromeres (Gray) Greene
Santa Rosa: Hoffmann s.n. Apr. 16, 1929 CAS# 168340!;
SBM.

Santa Cruz: Greene s.n. in Jul.-Aug. 1 886 ND-G#00 1494!;
Munz & Crow 11842 POM!; Raven & Smith 15235
RSA!; CAS; SBM.

Santa Catalina: Trask s.n. in Mar. 1 901 NY!; Thome 36872a
RSA!.

Cryptantha microstachys (Greene ex Gray) Greene
Santa Catalina: Knopf s.n. Mar. 20-Apr. 10, 1921 F!;
Millspaugh 4783 F!.

Cryptantha muricata (H. & A.) Nels. & Macbr. var. jonesii
(Gray) Jtn.

Santa Cruz: Hoffmann s.n. Apr. 22, 1932 POM#l 80293!;
SBM.

Cryptantha traskiae Jtn.

San Nicolas: Trask s.n. in Apr. 1901 LAM!; Howell 8223
CAS!; Wier & Beauchamp s.n. June 30, 1978 RSA!.
San Clemente: Trask 275 NY!; Raven 17271 RSA!; Pier-
son 3421 DS!

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 53

Harpagonella palmeri Gray

Santa Catalina: Johnson 1437 NY!; Thome 35873 RSA!;

Fosberg S4557 POM!; SBBG.

Guadalupe: Moran 6750 RSA!; Carlquist 463 RSA!.
Heliotropium curassavicum L. ssp. oculatum (Heller) Thome
San Miguel: Elmore 328 AHFH!; SBBG; SBM.

Santa Rosa: Dunkle 8470 LAM!, AHFH!; Thome et al.
48966 RSA!; SBBG.

Santa Cruz: Dunkle 8593 LAM!; Elmore 277 AHFH!; RSA-
POM; SBBG; SBM.

Anacapa: Dunkle 76 1 1 LAM!; Elmore 222 AHFH!; SBBG;
SBM.

San Nicolas: Trask s.n. in Apr. 1901 LAM!; Kanakoff s.n.
Apr. 12, 1940 LAM!; Dunkle 8319 LAM!; RSA-POM;
SBBG.

Santa Catalina: Fosberg 7166 LAM!, POM!; Thome &
Everett 34558 RSA!; Wolf 3602 RSA!; SBBG; SBM.
San Clemente: Murbarger 205 UC!.

Pectocarya linearis DC. ssp. ferocula (Jtn.) Thome

Santa Cruz: Fosberg 7623 LAM!; Clokey 5035 NY!; Raven
& Smith 15186 RSA!; SBBG; SBM.

Santa Catalina: Fosberg S4404 LAM!, NY, POM!; Grant
913 GH!; Thome & Everett 34670 RSA!; SBBG; SBM.
San Clemente: Trask 167 NY!; Munz 6690 POM!, GH!;

Thome 36101 RSA!.

Guadalupe: Carlquist 450 RSA!.

Pectocarya penicillata (H. & A.) A. DC.

Anacapa: SBM.

Pectocarya recurvata Jtn.

Guadalupe: Palmer 69a NY!; Anthony 237 GH!; Moran
6594 CAS!.

Plagiobothrys californicus (Gray) Greene var. ca/ifornicus
Guadalupe: Moran 20308 RSA!; Wiggins & Ernst 106 DS!.
Plagiobothrys californicus (Gray) Greene var. fulvescens Jtn.
San Miguel: SBM.

Santa Rosa: Munz & Crow 1 1705 POM!; Munz & Crow
11707 POM!.

Santa Cruz: Fosberg 7681 LAM!, LA!; SBBG; SBM.
Anacapa: SBBG; SBM.

Plagiobothrys californicus (Gray) Greene var. gracilis Jtn.
Santa Cruz: RSA-POM; SBM.

Anacapa: SBBG.

Santa Catalina: Fosberg S4379 LAM!; Dunkle 2061 LAM!;

Trask s.n. in Mar. 1901 NY!; RSA-POM; SBBG.

San Clemente: RSA-POM.

Plagiobothrys canescens Benth.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Hoffmann s.n. Apr. 10, 1931 LAM!; RSA-
POM; SBBG; SBM.

Santa Catalina: Fosberg 10648 LAM!, LA!; Grant &
Wheeler 6156 LAM!, NY!; Grant & Wheeler 6159 LAM!;
RSA-POM; SBBG; SBM.

San Clemente: Trask 210 NY!; RSA-POM.

Brassicaceae

Arabis glabra (L.) Bemh.

Santa Cruz: RSA-POM; SBM.

Arabis hoffmannii (Munz) Rollins
Santa Rosa: SBM.

Santa Cruz: RSA-POM; SBM
Athysanus pusillus (Hook.) Greene

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 LAM!; Fosberg
7630 LAM!; Fosberg 7691 LAM!; RSA-POM; SBBG;
SBM.

Santa Catalina: RSA-POM; SBBG.

Brassica geniculata (Desf.) J. Ball
San Miguel: SBBG.

Santa Rosa: SBBG.

Santa Cruz: Daily 500 SCIR!; SBBG.

San Nicolas: RSA-POM; SBBG.

Santa Catalina: Fosberg S5378 LAM!; RSA-POM; SBBG.
San Clemente: RSA-POM; SBBG.

Brassica kaber (DC.) L.C. Wheeler
Santa Cruz: SBM.

Santa Catalina: RSA-POM.

Brassica nigra (L.) Koch in Rohling
Santa Cruz: Hoffmann s.n. Jun. 15, 1930 LAM!; SBM.
San Nicolas: RSA-POM.

Santa Barbara: Dunkle 7445 LAM!; SBBG; SBM.

Santa Catalina: Fosberg S4720 LAM!; Dunkle 1918
AHFH!; Fosberg S4730 LAM!; NY; RSA-POM.

San Clemente: Trask 173 US!.

Guadalupe: Palmer 5 NY!.

Brassica rapa L. ssp. sylvestris (L.) Janchen
San Miguel: Dunkle 8423 AHFH!; SBM.

Santa Cruz: Hoffmann s.n. Apr. 10, 1930 LAM!; Dunkle
8637 LAM!; RSA-POM; SBM.

Santa Catalina: Detmers s.n. Apr. 14, 1929 USC!; Fosberg
SI 1872 LAM!; RSA-POM.

San Clemente: Raven 17618 RSA!.

Cakile edentula (Bigel.) Hook. var. edentula
San Miguel: Dunkle 8389 LAM!, AHFH!; SBBG; SBM.
Santa Rosa: SBM.

San Nicolas: SBBG; SBM.

Cakile maritima Scop. ssp. maritima
San Miguel: SBBG.

Santa Rosa: RSA-POM; SBBG.

Santa Cruz: RSA-POM; SBBG.

Anacapa: SBBG.

San Nicolas: Foreman 236 LA!; RSA-POM; SBBG.

Santa Catalina: RSA-POM; SBBG.

San Clemente: RSA-POM; SBBG.

Capsella bursa-pastoris (L.) Medic.

San Miguel: RSA-POM; SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Dunkle 8633 AHFH!; RSA-POM; SBBG; SBM.
Santa Catalina: Fosberg S46 1 5 LAM!; Dunkle 8633 LAM!;

Trask s.n. in Mar. 1901 NY!; RSA-POM; SBBG.

San Clemente: RSA-POM; SBBG.

Guadalupe: Wiggins & Ernst 66 DS!.

Cardamine californica (Nutt.) Greene
San Miguel: SBM.

Santa Rosa: SBM.

Santa Cruz: Fosberg 7583 LAM!; Moran 760 LAM!; Clo-
key 4944 LAM; RSA-POM; SBM.

54 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Santa Catalina: Trask s.n. in Mar. 1897 LAM!; Thorne &
Propst 42483 RSA!.

Cardamine oligosperma Nutt.

Santa Cruz: SBM.

Cardaria draba (L.) Desv.

Santa Cruz: Hoffmann 260 LAM!, LA!; RSA-POM; SBBG;
SBM.

Caulanthus inflatus Wats.

Santa Cruz: Elmore 453 AHFH!.

Caulanthus lasiophyllus (H. & A.) Pays.

San Miguel: RSA-POM; SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7702 LAM!; Abrams & Wiggins 169
NY!; RSA-POM; SBBG; SBM.

Anacapa: Moran 724 LAM!; SBBG; SBM.

Santa Barbara: SBBG.

Santa Catalina: Fosberg S4594 LAM!, NY!; Fosberg S4724
LAM!; Dunkle 2084 LAM!; RSA-POM; SBBG; SBM.
San Clemente: Trask 265 NY!; RSA-POM; SBBG.
Guadalupe: Palmer 880 ND-G!; Moran 1 3777 SD!; Moran
17360 SD!.

Descurainia pinnata (Walt.) Britt, ssp. menziesii (DC.) Detl.
Santa Rosa: Hoffmann s.n. May 10, 1932 SBM#12101!.
Santa Cruz: Fosberg 75 19 LAM!; Clokey 5987 NY!; Abrams
& Wiggins 56 CAS!, DS!; RSA-POM; SBBG; SBM.
Anacapa: SBBG.

Santa Catalina: Dunkle 2092 AHFH!; Trask s.n. in Mar.

1901 NY!; RSA-POM; SBBG; SBM.

San Clemente: RSA-POM; SBBG.

Guadalupe: Palmer 6 NY!; Moran 5658 DS!.

Dithyrea maritima A. Davids.

San Miguel: SBM.

San Nicolas: Trask 29 NY!, US!; Trask s.n. in Apr. 1897
MO!; Raven & Thompson 20706 LA!; RSA-POM;
SBBG.

Draba cunefolia Nutt, ex T. & G. var. integrifolia Wats.

Santa Catalina: Trask s.n. in Apr. 1902 LAM!, NY!.
Erysimum ammophilum Heller

Santa Rosa: Munz & Crow 1 1757 LA!.

Erysimum cheiri (L.) Crantz

Santa Catalina: Thome & Everett 33447 RSA!.
Erysimum insulare Greene

San Miguel: Dunkle 8384 LAM!; Munz & Norris 1 1787
LA!; Greene s.n. in Sep. 1886 CAS!; RSA-POM; SBBG;
SBM.

Santa Rosa: Moran 797 LAM!; Moran s.n. Apr. 4, 1941
LAM!; Dunkle 8479 AHFH!; RSA-POM; SBBG.
Anacapa: SBBG.

Erysimum moranii Roll.

Guadalupe: Rempel 759-37 LAM!; Moran 15116 RSA!;
Moran 18169 RSA!.

Hutchinsia procumbens (L.) Desv.

San Miguel: Dunkle 8403 LAM!, AHFH!; Fosberg S2083
NY!; RSA-POM.

Santa Rosa: Munz & Crow 1 1754 LA!; RSA-POM.

Santa Cruz: RSA-POM.

Anacapa: SBBG.

San Nicolas: Dunkle 8320 LAM!; Trask 98 NY!; Raven
& Thompson 20685 DS!.

Santa Barbara: SBBG.

Guadalupe: Howell 8330 CAS!.

Lepidium lasiocarpum Nutt, ex T. & G. var. lasiocarpum
San Miguel: Dunkle 8422 LAM!.

Santa Rosa: RSA-POM.

Santa Cruz: SBBG.

San Nicolas: Trask s.n. in Apr. 1901 LAM!; Dunkle 8320a
LAM!; Kanakoff s.n. Apr. 13, 1940 LAM!; RSA-POM;
SBBG.

Santa Catalina: Millspaugh 4791 F!; Millspaugh 4614 F!.
San Clemente: RSA-POM; SBBG.

Guadalupe: Moran 6718 RSA!; Moran 2889 DS!; Moran
5696 DS!.

Lepidium lasiocarpum Nutt, ex T. & G. var. latifolium C. L.
Hitchc.

Guadalupe: Palmer 8 NY!; Palmer 841 NY!, US!; Mason
1516 US!.

Lepidium latipes Hook.

Santa Cruz: RSA-POM; SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 F!, NY!; Mills-
paugh 4749 F!.

San Clemente: RSA-POM.

Lepidium nitidum Nutt. var. nitidum
Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7526 LAM!, LA!; Fosberg 7587 LAM!;

RSA-POM; SBBG; SBM.

Anacapa: SBBG; SBM.

San Nicolas: Foreman 227 US!; SBM.

Santa Barbara: Dunkle 74 1 3 LAM!; Bryan, Dr. & Mrs. s.n.
LAM!; RSA-POM; SBBG.

Santa Catalina: Fosberg S4391 LAM!; Dunkle 2071
AHFH!; Trask s.n. in Mar. 1901 NY!; RSA-POM; SBBG.
San Clemente: Elmore 401 AHFH!; Trask 347 NY!; RSA-
POM; SBBG.

Guadalupe: Wiggins & Ernst 1 10 DS!; Moran 5677 SD!;
Moran 12047 SD!.

Lepidium oblongum Small
San Miguel: RSA-POM.

Santa Cruz: RSA-POM.

Anacapa: SBBG.

San Nicolas: Trask 28 GH!; SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 NY!.

San Clemente: SBBG.

Guadalupe: Moran 2878 DS!; Palmer 897 GH!, US!; Rose
16006 GH!, NY!, US!.

Lepidium virginicum L. var. pubescens (Greene) Thell.
Santa Catalina: Dunkle 2454 AHFH!; RSA-POM; SBBG.
San Clemente: SBBG.

Lepidium virginicum L. var. robinsonii (Thell.) C.L. Hitchc.
Santa Cruz: RSA-POM; SBM.

San Nicolas: Trask s.n. in Apr. 1901 LAM!; Kanakoff s.n.
Apr. 12, 1940 LAM!; Dunkle 8320 MO!; RSA-POM;
SBBG.

Lobularia maritima (L.) Desv.

San Nicolas: Newman 103 Pacif. Missile Test Center Pt.
Mugu!.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 55

Santa Catalina: Fosberg 4483 LAM!; Lister & Powell s.n.
Apr. 2, 1928 USC!; Trask s.n. in 1901 NY!; RSA-POM;
SBBG.

Nasturtium officinale R. Br.

San Miguel: SBBG.

Santa Cruz: RSA-POM; SBM.

Santa Catalina: Fosberg S4748 LAM!; Dunkle 1925
AHFH!; RSA-POM; SBBG; SBM.

Raphanus raphanistrum L.

Santa Rosa: RSA-POM; SBBG.

Santa Catalina: RSA-POM; SBBG.

San Clemente: RSA-POM; SBBG.

Raphanus sativus L.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Fosberg
S4663 LAM!; RSA-POM; SBBG.

San Clemente: RSA-POM; SBBG.

Sibara flifolia (Greene) Greene
Santa Cruz: SBM.

Santa Catalina: Trask s.n. in May 1901 LAM!.
Sisymbrium altissimum L.

Santa Catalina: Fosberg 4854 LAM!; Fosberg S5385 LAM!.
Sisymbrium irio L.

Anacapa: SBBG.

Santa Catalina: RSA-POM; SBBG.

San Clemente: RSA-POM; SBBG.

Guadalupe: Howell 8276 CAS!.

Sisymbrium officinale (L.) Scop.

Santa Cruz: Daily 336 SCIR!; SBBG; SBM.

Santa Catalina: Fosberg S4465 LAM!; Trask 303 US!, NY!;
Trask s.n. in Apr. 1900 NY!.

Sisymbrium orientale L.

Santa Catalina: RSA-POM; SBBG.

Thysanocarpus curvipes Hook. ssp. curvipes
Santa Cruz: Hoffmann s.n. Apr. 12, 1931 LA!.
Thysanocarpus curvipes Hook. var. elegans (F. & M.) Rob.
in Gray

Santa Catalina: RSA-POM.

Thysanocarpus erectus Wats.

Guadalupe: Anthony 244 SD!.

Thysanocarpus laciniatus Nutt, ex T. & G. var. conchulifer-
us (Greene) Jeps.

Santa Cruz: Moran 755 LAM!; Williams s.n. Mar. 23, 1941
AHFH!; Munz & Crow 1 1848 LA!; SBM.
Thysanocarpus laciniatus Nutt, ex T. & G. var. crenatus
(Nutt.) Brewer
Santa Rosa: SBM.

Santa Cruz: SBM.

Thysanocarpus laciniatus Nutt, ex T. & G. var. laciniatus
Santa Cruz: Moran 750 LAM!; Williams s.n. Mar. 23, 1941
AHFH!; Hoffmann s.n. Apr. 10, 1931 LAM!; LA; RSA-
POM.

Santa Catalina: Fosberg S4309 LAM!; Dunkle 2123
AHFH!; Detmers s.n. Apr. 13, 1929 USC!.

San Clemente: Dunkle 7308 LAM!, AHFH!.
Thysanocarpus laciniatus Nutt, ex T. & G. var. ramosus
(Greene) Munz

Santa Rosa: SBM.

Santa Cruz: Fosberg 7626 LAM!, LA!; SBM.
Tropidocarpum gracile Hook.

Santa Catalina: Trask s.n. in 1900 LAM!; Dunkle 2064
AHFH!; SBBG.

San Clemente: RSA-POM.

Cactaceae

Bergerocactus emoryi (Engelm.) Britt. & Rose
Santa Catalina: RSA-POM; SBBG; SBM.

San Clemente: Dunkle 7218 AHFH!; RSA-POM; SBBG;
SBM.

Mammillaria blossfeldiana Bodeker var. shurliana Gates
Guadalupe: Moran 6708A SD!; Moran 17413 SD!; Moran
15123 SD!.

Opuntia ficus-indica (L.) Mill.

Santa Catalina: Millspaugh 4523 F!.

Opuntia littoralis (Engelm.) Ckll.

San Miguel: SBM.

Santa Rosa: SBM.

Santa Cruz: Elmore 279 AHFH!; RSA-POM; SBBG; SBM.
Anacapa: SBBG; SBM.

San Nicolas: SBBG.

Santa Barbara: Philbrick 0-628 SBBG!.

Santa Catalina: Millspaugh 4525 F!.

San Clemente: Blakley 5248 RSA!.

Opuntia oricola Philbrick
San Miguel: SBBG.

Anacapa: Elmore 245 LAM!; SBBG.

San Nicolas: Raven & Thompson 20790 RSA!; SBBG.
Santa Barbara: Blakley 5713 RSA!; SBBG.

Santa Catalina: Fosberg S4892 LAM!, F!.

San Clemente: Elmore 382 LAM!; Dunkle 7217 AHFH!.
Opuntia prolifera Engelm.

Santa Rosa: SBBG.

Santa Cruz: SBBG.

Anacapa: SBBG; SBM.

San Nicolas: SBBG.

Santa Barbara: SBBG.

Santa Catalina: Fosberg S4733 LAM!.

San Clemente: RSA-POM; SBBG.

Guadalupe: Moran 2617 SD!.

Callitrichaceae

Callitriche longipedunculata Morong
San Clemente: Thome 42802 RSA!.

Callitriche marginata Torr. var. marginal a
Santa Catalina: Thome 37697 RSA!; Thome 36088 RSA!.
San Clemente: Raven 17328 RSA!.

Campanulaceae

Githopsis diffusa Gray ssp. diffusa
Santa Cruz: Hoffmann s.n. Apr. 21, 1932 POM!; Hoffmann
s.n. May 23, 1932 POM!; Raven & Smith 15236 RSA!.
Guadalupe: Palmer 53 CM!; Moran 27271 SD!. (GU ma-
terial is G. diffusa Gray var. guadalupensis Morin (Mo-
rin, N. Systematic Botany 8(4):436-468. 1983.)

Lobelia erinus L.

Santa Cruz: SBBG.

56 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Triodanis bi flora (R. & P.) Greene
Santa Rosa: SBM.

Santa Cruz: Hoffmann s.n. Jun. 13, 1930 POM!; Raven &
Smith 15150 RSA!; SBBG; SBM.

Santa Catalina: FosbergS481 1 LAM!; Fosberg4781 LAM!;
Thorne 36334 RSA!.

Guadalupe: Moran 6611 SD!; Moran 6641 SD!; Moran
13810 SD!.

Capparaceae

Cleome isomeris Greene

Santa Rosa: Moran 786 LAM!; Dunkle 8441 LAM!,
AHFH!; RSA-POM; SBBG; SBM.

Santa Catalina: Grant 615 LAM!; Trask s.n. in Jul. 1898
US!; Carlson s.n. May 1 0, 1 9 1 8 US!; RSA-POM; SBBG;
SBM.

San Clemente: SBBG.

Caprifoliaceae

Lonicera hispidula (Lindl.) Dougl. ex T. & G. var. vacillans
Gray

Santa Cruz: Hoffmann s.n. Jul. 1, 1930 LAM!; Hoffmann
s.n. Sep. 10, 1931 LAM!; RSA-POM; SBBG; SBM.
Santa Catalina: Fosberg S485 1 LAM!, NY!; Knopf 1 39 F!;

Trask s.n. in Jun. 1895 US!; LA; RSA-POM; SBBG.
San Clemente: Trask 21 US!; Trask 197 NY!, US!; RSA-
POM.

Lonicera subspicata H. & A. var. johnstonii Keck
Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: RSA-POM; SBBG.

Sambucus mexicana Presl ex DC.

Santa Rosa: Dunkle 8508 LAM!; RSA-POM; SBM.

Santa Cruz: Hoffmann s.n. Sep. 21, 1930 LAM!; Hoffmann
s.n. Apr. 12, 1931 LAM!; RSA-POM; SBBG; SBM.
Santa Catalina: Fosberg S4289 LAM!; Dunkle 1984
AHFH!; Detmers s.n. Apr. 14, 1929 USC!; RSA-POM;
SBBG; SBM.

San Clemente: House & Grumbles s.n. Aug. 5-13, 1930
USC!; RSA-POM; SBBG.

Symphoricarpos mollis Nutt, in T. & G.

Santa Rosa: SBM.

Santa Cruz: Dunkle 8557 LAM!; Munz & Crow 11550
POM!, LA!; Raven & Smith 15180 RSA!; SBBG; SBM.
Santa Catalina: Fosberg S4300 LAM!; Dunkle 2135 LAM!;
Detmers s.n. in Spring 1928 USC!; RSA-POM; SBBG.
Caryophyllaceae

Arenaria douglasii Fenzl ex T. & G.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: SBBG; SBM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Fosberg
S4721 LAM!; RSA-POM!.

San Clemente: RSA-POM.

Arenaria serpyllifolia L.

Santa Cruz: RSA-POM.

Cardionema ramosissima (Weinm.) Nels. & Macbr.

San Miguel: SBBG; SBM.

Santa Rosa: Dunkle 8466 LAM!, AHFH!; Dunn, N. s.n.

May 24, 1931 LA!; RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7670 LAM!, LA!; RSA-POM; SBBG;
SBM.

Cerastium glomeratum Thuill.

San Miguel: RSA-POM; SBBG; SBM.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: Fosberg 7669 LAM!; RSA-POM; SBBG; SBM.
Santa Catalina: RSA-POM; SBBG.

San Clemente: RSA-POM; SBBG.

Guadalupe: Moran 17304 RSA!.

Herniaria cinerea DC.

Guadalupe: Wiggins & Ernst 195 DS!.

Polycarpon depression Nutt.

Santa Cruz: SBM.

Santa Catalina: Grant s.n. in Apr. 1901 LAM!; Grant 1023
US!; Trask s.n. in Jun. 1897 US!; SBBG.

Sagina decumbens (Ell.) T. & G. ssp. occidentalis (Wats.)
Crow

San Miguel: SBM.

Santa Rosa: SBM.

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 LAM!; Wolf 2909
DS!; Hoffmann s.n. Jun. 15, 1930 CAS#191796!; RSA-
POM; SBM.

Santa Catalina: Fosberg S4331 LAM!.

Silene antirrhina L.

Santa Rosa: SBBG; SBM.

Santa Cruz: RSA-POM; SBM.

Santa Catalina: Fosberg S4381 LAM!; Trask s.n. in May
1 897 US!; Trask s.n. in Mar. 1 898 MO!; NY; RSA-POM;
SBBG.

San Clemente: Trask 175 NY!, US!; RSA-POM; SBBG.
Guadalupe: Gander 8999 SD!.

Silene ga/lica L.

San Miguel: SBBG; SBM.

Santa Rosa: Dunkle 8528 LAM!; Dunn, N. s.n. May 24,
1931 LA!; Dunn, D. 1313 LA!; RSA-POM; SBM.
Santa Cruz: Clokey 4933 LAM!; Clokey 4934 LAM!; El-
more 456 AHFH!; RSA-POM; SBM.

Anacapa: Dunkle 7662 LAM!; SBBG; SBM.

San Nicolas: Trask 25 MO!.

Santa Barbara: Dunkle 7645 LAM!; Dunkle 8143 LAM!;
RSA-POM; SBBG; SBM.

Santa Catalina: Moran 705 LAM!; Dunkle 1827 AHFH!;

Detmers s.n. Apr. 13, 1929 USC!; RSA-POM.

San Clemente: Dunkle 7305 LAM!; Dunkle 7259 LAM!;
RSA-POM; SBBG.

Guadalupe: Palmer 1 1 NY!; Palmer 837 NY!; Lindsay
43072 RSA!.

Silene laciniata Cav. ssp. major Hitchc. & Maguire
San Miguel: RSA-POM; SBM.

Santa Rosa: Moran 809 LAM!, MO!, NY!; Elmore 206
AHFH!; Epling & Erickson s.n. Aug. 8, 1937 LA!; RSA-
POM; SBM.

Santa Cruz: Clokey 493 1 LAM!, NY!; Eastwood 6408 NY!,
US!; Johnstone s.n. USC!; RSA-POM; SBM.

Anacapa: Dunkle 7654 LAM!; Elmore 239 AHFH!; Ellison
s.n. May 12-15, 1929 LA!; RSA-POM; SBBG; SBM.
San Clemente: Trask 36 US!.

Silene multinervia Wats.

Santa Cruz: RSA-POM; SBM.

Santa Catalina: Trask s.n. in Mar. 1900 MO!, US!.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 57

S per gill a arvensis L.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Catalina: RSA-POM; SBBG.

Spergularia bocconii (Scheele) Foucaud
Santa Cruz: RSA-POM; SBBG.

Santa Catalina: Davidson, A. s.n. Jun. 25, 1891 LAM!;

Fosberg S4952 LAM!; Wolf 3578 LAM!; RSA-POM.
San Clemente: RSA-POM; SBBG.

Spergularia macrotheca (Homem.) Heynh. ssp. macrotheca
San Miguel: Dunkle 8398 LAM!; RSA-POM; SBM.

Santa Rosa: Dunkle 8484 LAM!; Elmore 1 89 AFIFFI!; RSA-
POM; SBM.

Santa Cruz: Fosberg 7534 LAM!; Dunkle 8576 LAM!,
AHFH!; Elmore s.n. Apr. 17, 1936 USC!; RSA-POM;
SBM.

Anacapa: Gustafson s.n. Apr. 9, 1973 LAM!; Johnstone
s.n. Jun. 25, 1932 USC!; SBBG; SBM.

San Nicolas: Kanakoff s.n. Apr. 12, 1940 LAM!; Dunkle
8321 LAM!; Raven & Thompson 20691 LA!; RSA-POM;
SBM.

Santa Barbara: Dunkle 8110 AHFH!; Bryan, Dr. & Mrs.

s.n. Jul. 14, 1922 LAM!; Dunkle 8130 LAM!; SBBG.
Santa Catalina: Moran 664 LAM!; Fosberg S4466 LAM!;

Dunkle 1853 AHFH!; RSA-POM; SBM.

San Clemente: Dunkle 7205 LAM!; Dunkle 7272 LAM!;
Elmore s.n. Nov. 25, 1939 AHFH!; RSA-POM; SBBG;
SBM.

Guadalupe: Rempel 758-37 LAM!; Moran 6126 RSA!;
Moran 17417 RSA!.

Spergularia marina (L.) Griseb.

Santa Rosa: RSA-POM; SBBG.

Santa Cruz: RSA-POM; SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Fosberg
S4332 LAM!; Dunkle 1737 AHFH!; RSA-POM.

San Clemente: Murbarger 1 12 UC!.

Guadalupe: Moran 6768 RSA!.

Spergularia villosa (Pers.) Camb.

Santa Rosa: RSA-POM; SBBG.

Santa Catalina: RSA-POM; SBBG.

San Clemente: RSA.

Stellaria media (L.) Vill.

San Miguel: SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Clokey 4928 LAM!; Clokey 4929 LAM!; Dun-
kle 8624 LAM!; RSA-POM; SBBG; SBM.

Santa Catalina: Moran 706 LAM!; Fosberg S4644 LAM!;

Dunkle 2144 AHFH!; RSA-POM; SBBG.

San Clemente: RSA-POM; SBBG.

Stellaria nitens Nutt.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: Fosberg 7575 LAM!; Fosberg 7600 LAM!;

Fosberg s.n. Mar. 6, 1932 LA!; RSA-POM; SBBG; SBM.
Santa Catalina: Dunkle 2089 AHFH!; McClatchie s.n. in
Sep. 1893 NY!; Trask s.n. in Mar. 1901 NY!.
Guadalupe: Palmer 13 NY!; Moran 18388 SD!; Moran
25386 SD!.

Chenopodiaceae

Aphanisma blitoides Nutt, ex Moq. in DC.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: RSA-POM.

Anacapa: Hoffmann s.n. Mar. 21, 1932 SBM!.

San Nicolas: Trask 21 MO!.

Santa Barbara: Dunkle 7416 LAM!, DS!, NY!; Dunkle
7459 LAM!, AHFH!; RSA-POM; SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Trask s.n.
in May 1900 NY!; Grant & Wheeler s.n. Apr. 21-26,
1904 DS!; RSA-POM.

San Clemente: Trask s.n. in Apr. 1898 US!; Trask 212
NY!, US!; Blakley 6359 DS!; RSA-POM.

Guadalupe: Moran 5627 CAS!, DS!, RSA!; Moran 5656
DS!, RSA!; Wiggins & Ernst 175 DS!.

Atriplex argentea Nutt. ssp. expansa (Wats.) Hall & Clem.
Santa Rosa: RSA-POM; SBM.

Santa Cruz: RSA-POM; SBM.

Santa Catalina: RSA-POM; SBM.

San Clemente: RSA-POM; SBBG.

Atriplex barclayana (Benth.) D. Dietr. ssp. dilitata (Greene)
Hall & Clem.

Guadalupe: Rempel 759-37 Jul. 19, 1937 LAM!.

Atriplex barclayana (Benth.) D. Dietr. ssp. palmeri (Wats.)
Hall & Clem.

Guadalupe: Greene s.n. in 1885 US!; Palmer 863 US!;
Moran 5622 RSA!.

Atriplex californica Moq. in DC.

San Miguel: Dunkle 8370 LAM!, Dunkle 8370 AHFH!;
RSA-POM; SBM.

Santa Rosa: Dunkle 8485 LAM!; RSA-POM; SBM.

Santa Cruz: Fosberg 7704 LAM!; Fletcher s.n. LAM!;

Clokey 4920 LAM!; RSA-POM; SBM.

Anacapa: Dunkle 7604 LAM!; Dunn, N. s.n. May 14, 1932
LA!; RSA-POM; SBM.

San Nicolas: Dunkle 8317 LAM!; Kanakoff s.n. Apr. 24,
1940 LAM!; Dunkle 8343 LAM!; RSA-POM; SBBG.
Santa Barbara: RSA-POM; SBM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Dunkle
7410 LAM!; Fosberg S4889 LAM!; RSA-POM; SBM.
San Clemente: Dunkle 7275 LAM!; Johnstone s.n. Sep. 5,
1926 USC!; RSA-POM.

Guadalupe: Moran 15126 RSA!.

Atriplex coulteri (Moq.) D. Dietr.

San Miguel: SBBG; SBM.

Santa Rosa: Hoffmann s.n. Jun. 13, 1930 POM!; RSA-
POM; SBM.

Santa Cruz: RSA-POM; SBM.

Anacapa: SBM.

San Nicolas: RSA-POM.

Santa Catalina: Fosberg S4648 LAM!; Fosberg S4669
LAM!; Fosberg S4730 LAM!; RSA-POM; SBBG.

San Clemente: RSA-POM.

Atriplex lentiformis (Torr.) Wats. ssp. breweri (Wats.) Hall &
Clem.

Santa Cruz: SBBG; SBM.

Anacapa: Dunkle 7640 LAM!, NY!; RSA-POM; SBBG;
SBM.

58 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Santa Catalina: Trask s.n. in Jun. 1897 US!; Trask s.n. in
Jun. 1898 US!; Millspaugh 5486 F!.

San Clemente: Trask s.n. Oct. 19, 1902 US!; RSA-POM;
SBBG; SBM.

Atriplex leucophylla (Moq. in DC.) D. Dietr.

San Miguel: RSA-POM; SBM.

Santa Rosa: RSA-POM.

Santa Cruz: RSA-POM; SBM.

San Nicolas: Kanakoff s.n. Apr. 13, 1940 LAM!; Dunkle
8340 LAM!; Trask s.n. Apr. 1901 LAM!; RSA-POM;
SBM.

Santa Catalina: Fosberg S4894 LAM!; Trask s.n. in Aug.

1902 LAM!; Dunkle 1852 AHFH!; RSA-POM.

San Clemente: Trask s.n. Nov. 22, 1 922 US#6 1 7893!; RSA-
POM.

Atriplex pacifica Nels.

Santa Cruz: Greene s.n. in Jul.-Aug. 1 886 ND-G#01 5274!,
US!.

Anacapa: SBBG; SBM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!, US!; Grant
& Wheeler s.n. Apr. 21-26, 1 904 LAM!; McClatchie s.n.
Sep. 13, 1893 NY!; SBBG.

San Clemente: Trask s.n. in May 1901 NY!; Wooton s.n.
Apr. 26, 1912 US!; RSA-POM.

Atriplex patula L. ssp. hastata (L.) Hall & Clem.

San Miguel: RSA-POM; SBBG; SBM.

Santa Cruz: RSA-POM; SBBG.

San Nicolas: SBBG.

Santa Catalina: Dunkle 2440 LAM!, AHFH!; RSA-POM;
SBBG.

Atriplex rosea L.

Santa Catalina: RSA-POM.

Atriplex semibaccata R. Br.

San Miguel: Dunkle 8380 AHFH!; Dunkle 8418 LAM!;

RSA-POM; SBM.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: RSA-POM.

Anacapa: SBBG; SBM.

San Nicolas: Dunkle 8304 LAM!, AHFH!; Kanakoff s.n.
Apr. 13, 1940 LAM!; Dunkle s.n. Nov. 24, 1940 LAM!;
RSA-POM; SBM.

Santa Barbara: Dunkle 8123 AHFH!; Bryan, Dr. & Mrs.

s.n. Jul. 14, 1922 LAM!; RSA-POM; SBM.

Santa Catalina: Dunkle 1817 AHFH!; Spalding s.n. in 1 925
USC!; Fosberg S4569 LAM!; RSA-POM; SBM.

San Clemente: Elmore 419 LAM!; Dunkle 8123 LAM!;
House & Grumbles s.n. Aug. 5-13, 1930 USC!; RSA-
POM; SBM.

Atriplex serenana A. Nels. var. serenana

Santa Catalina: Moxley s.n. Sep. 19, 1925 USC!; Nuttall
303 F!; Eastwood 6529 US!.

Atriplex watsonii A. Nels. in Abrams

Santa Cruz: Ferren 1907a UCSB!; Ferren 1907b UCSB!.
San Nicolas: Dunkle 83 1 7 AHFH!; Trask s.n. in Apr. 1897
US!; RSA-POM; SBBG.

Santa Catalina: Fosberg S4925 LAM!; Fosberg S4928 NY!;
RSA-POM.

San Clemente: Dunkle 7264 LAM!; AHFH!; Dunkle 7274

AHFH!, LAM!; Trask s.n. in Oct. 1 902 US!; RSA-POM;
SBBG.

Bassia hyssopifolia (Pall.) Kuntze
Santa Catalina: RSA-POM; SBBG.

San Clemente: RSA-POM.

Beta vulgaris L. ssp. maritima (L.) Arcang.

San Miguel: SBBG.

Anacapa: SBBG.

Santa Catalina: RSA-POM.

San Clemente: RSA-POM.

Chenopodium album L.

Santa Cruz: Clokey 4917 LAM!; Howell 6237 CAS!.
Chenopodium ambrosioides L. var. ambrosioides
Santa Rosa: Dunkle 8465 LAM!, AHFH!; Dunkle 8514
LAM!, NY!; Dunkle 8527 LAM!; RSA-POM; SBBG;
SBM.

Santa Cruz: Dunkle 8645 LAM!, AHFH!; Clokey 4916
NY!; RSA-POM; SBBG; SBM.

Santa Catalina: RSA-POM; SBBG.

Chenopodium berlandieri Moq. var. sinuatum (J. Murr.) H.A.
Wahl.

Santa Rosa: RSA-POM.

Santa Cruz: Hoffmann s.n. Apr. 10, 1931 LAM!; RSA-
POM; SBBG; SBM.

Anacapa: RSA-POM; SBBG.

Santa Catalina: Fosberg S5376 LAM!; Fosberg S4508
LAM!; Dunkle 2005 AHFH!; RSA-POM.
Chenopodium californicum (Wats.) Wats.

San Miguel: RSA-POM; SBM.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: Fosberg 7554 LAM!; Clokey 49 1 5 LAM!; RSA-
POM; SBM.

Anacapa: Bond 351 SBM!; SBBG; SBM.

San Nicolas: RSA-POM.

Santa Barbara: Bond 392 SBM!; Dunkle 7453 LAM!; Dun-
kle 8116 LAM!.

Santa Catalina: Templeton 1 1382 LAM!; Fosberg S4283
LAM!, SBM!; Lewis s.n. Mar. 24, 1937 LA!; RSA-POM.
San Clemente: Moran 578 LAM!; RSA-POM.
Chenopodium multifidum L.

Santa Rosa: RSA-POM.

San Nicolas: Kanakoff s.n. Apr. 18, 1940 LAM!.
Chenopodium murale L.

San Miguel: Dunkle 8392 AHFH!; RSA-POM; SBM.
Santa Rosa: Dunkle 8468 LAM!, AHFH!; Elmore 178
AHFH!; RSA-POM; SBM.

Santa Cruz: Clokey 49 1 3 LAM!, NY!; Clokey 4914 LAM!,
NY!; Dunkle 8592 AHFH!; RSA-POM; SBM.
Anacapa: Dunkle 7610 LAM!; SBBG; SBM.

San Nicolas: Dunkle 8322 LAM!, AHFH!; Trask s.n. in
Apr. 1897 US!; RSA-POM.

Santa Barbara: Dunkle 8 1 00 LAM!, NY!; Bond 39 1 SBM!;

Philbrick B68-98 US!; RSA-POM.

Santa Catalina: Trask s.n. in 1900 LAM!, NY!; Fosberg
7162 LAM!; Smith 5071 US!; RSA-POM.

San Clemente: Dunkle 7222 LAM!; Trask 38 US!; Fosberg
55249 US!; RSA-POM.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 59

Guadalupe: Franceschi s.n. in 1893 US!; Rose 16037 NY!,
US!; Mason 1520 CAS!; MO.

Monolepis nuttalliana (Schult.) Greene
San Miguel: RSA-POM; SBM.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: SBM.

San Clemente: RSA-POM; SBBG.

Salicornia subterminalis Parish
San Miguel: SBBG.

Santa Rosa: RSA-POM; SBBG.

Santa Cruz: RSA-POM.

Santa Catalina: Dunkle 1897 AHFH!; Fosberg S4886
LAM!; Fosberg S4888 LAM!; RSA-POM; SBM.

San Clemente: House & Grumbles s.n. Aug. 5-13, 1930
USC!; RSA-POM; SBBG.

Salicornia virginica L.

San Miguel: SBBG.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: RSA-POM.

Anacapa: Dunkle 7652 LAM!; SBBG.

San Nicolas: Trask s.n. in Apr. 1901 LAM!; KanakofF s.n.

Apr. 19, 1940 LAM!; RSA-POM.

Santa Catalina: Fosberg S4878 LAM!; Fosberg S5426
LAM!; Williamsen s.n. in Aug. 1924 USC!; RSA-POM.
San Clemente: Dunkle 7330 LAM!; Elmore 402 AHFH!;
RSA-POM; SBBG.

Salsola iberica Sennen & Pau
Santa Cruz: Dunkle 8653 LAM!; RSA-POM.

San Nicolas: RSA-POM.

Santa Catalina: Fosberg S4967 LAM!.

San Clemente: RSA-POM.

Suaeda californica Wats.

San Miguel: RSA-POM; SBBG; SBM.

Santa Rosa: Munz & Crow 1 1565 LA!; RSA-POM; SBBG;
SBM.

Santa Cruz: Clokey 4921 LAM!, US!; Clokey 4922 LAM!,
US!; Fosberg 7706 LAM!; RSA-POM; SBM.

Anacapa: Dunkle 7602 LAM!; Howell 3811 CAS!; RSA-
POM; SBM.

San Nicolas: Trask 1 9 NY!; Foreman 5 1 US!; Dunkle 8313
LAM!, AHFH!; RSA-POM; SBM.

Santa Barbara: Philbrick& Benedict B66-417 US!; Dunkle
8114 LAM!, AHFH!; Elmore 302 AHFH!; RSA-POM;
SBM.

Santa Catalina: Trask s.n. in Mar. 1901 F!, US!; Fosberg
4913 LAM!; Nuttall 225 F!; RSA-POM; SBM.

San Clemente: Dunkle 7265 LAM!, AHFH!; Trask 36 US!;
House & Grumbles s.n. Aug. 5-13, 1930 USC!; RSA-
POM; SBBG.

Guadalupe: Moran 13761 LAM!, RSA!; Rempel 758-37
LAM!; Palmer 870 NY!; CAS.

Cistaceae

Helianthemum greenei Rob.

Santa Rosa: Epling & Erickson s.n. Aug. 8, 1937 LA!;

Dunn, N. s.n. May 24, 1931 LA!.

Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: RSA-POM; SBBG.

Helianthemum scoparium Nutt.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Fosberg 8157 LA!; RSA-POM; SBBG;
SBM.

Convolvulaceae

Calystegia macrostegia (Greene) Brummitt ssp. amplissima
Brum mitt

San Nicolas: Dunkle 8344 LAM!, AHFH!; Trask 53 US!;

Foreman 211 LA!, US!; RSA-POM; SBBG; SBM.
Santa Barbara: Dunkle 8138 LAM!, AHFH!; Elmore 313
AHFH!; Dunkle 8317 AHFH!; RSA-POM; SBBG; SBM.
San Clemente: Munz 6622 LAM!; Wiggins 11964 DS!;
Raven 18030 RSA!; SBBG; US.

Calystegia macrostegia (Greene) Brummitt ssp. cyclostegia
(House) Brummitt

Santa Catalina: Fosberg 8138 LA!; RSA-POM; SBM.
Calystegia macrostegia (Greene) Brummitt ssp. intermedia
(Abrams) Brummitt

Santa Catalina: Fosberg S4335 LAM!; Millspaugh 4657 F!;
RSA-POM.

Calystegia macrostegia (Greene) Brummitt ssp. macrostegia
San Miguel: Dunkle 8386 LAM!, AHFH!; SBBG; SBM.
Santa Rosa: Dunkle 8457 LAM!; RSA-POM; SBBG; SBM.
Santa Cruz: Greene s.n. in Jul.-Aug. 1886 US!; Fosberg
7703 LAM!; Dunkle 8541 AHFH!; LA; RSA-POM;
SBBG; SBM.

Anacapa: Moran 720 LAM!; Dunkle 7626 AHFH!; Ellison
s.n. May 12-15, 1929 LA!; RSA-POM; SBBG; SBM.
Santa Catalina: Trask s.n. in May 1901 LAM!; Dunkle
1769 AHFH!; Lister & Powell s.n. Apr. 3, 1928 USC!.
Guadalupe: Rempel 759-37 LAM!; Moran 2636 RSA!;
Carlquist 471 RSA!.

Calystegia soldanella (L.) R. Br.

San Miguel: RSA-POM; SBBG; SBM.

Santa Rosa: RSA-POM; SBBG.

Santa Cruz: RSA-POM; SBBG; SBM.

San Clemente: Murbarger 209 UC!.

Convolvulus arvensis L.

Santa Rosa: RSA-POM.

Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Dunkle 1947 AHFH!; Fosberg S4745
LAM!; RSA-POM; SBBG; SBM.

Convolvulus simulans L.

San Clemente: SBBG.

Cressa truxillensis HBK. var. vallicola (Heller) Munz
San Miguel: SBBG.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: RSA-POM; SBM.

Santa Catalina: Fosberg S4880 LAM!, NY!; Trask s.n. Jun.
1900 NY!; Dunkle 2147 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

San Clemente: Trask 184 NY!; RSA-POM; SBBG.
Cuscuta californica H. & A.

Santa Rosa: RSA-POM; SBM.

San Clemente: Trask 187 US!.

Cuscuta ceanothi Behr

Santa Cruz: Clark s.n. Mar. 27, 1980 SCIR!; SBBG.

60 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Cuscuta corymbosa R. & P. var. grandiflora Engelm.

Guadalupe: Moran 17420 SD!.

Cuscuta occidentalis Millsp.

Santa Catalina: Nuttall 902 F!; Nuttall 272 F!; RSA-POM;
SBBG.

Cuscuta salina Engelm. var. salina
Santa Rosa: RSA-POM.

Santa Cruz: RSA-POM; SBBG.

Anacapa: SBBG.

Dichondra occidentalis House
San Miguel: SBBG.

Santa Rosa: Munz & Crow 1 1574 POM!; Hoffmann s.n.
Apr. 18, 1932 POM!; SBM.

Santa Cruz: Wolf 2841 POM!; RSA-POM; SBBG; SBM.
Santa Catalina: RSA-POM; SBBG.

Ipomoea cairica (L.) Sweet
Santa Catalina: Spalding s.n. Sep. 19, 1925 USC!.
Ipomoea nil (L.) Roth

Santa Catalina: Millspaugh 4541 F!.

Comaceae

Cornus glabrata Benth.

Santa Catalina: Trask s.n. in Jun. 1901 LAM!; Dunkle
2032 LAM!; Knopf 126 F!; RSA-POM; SBM.
Crassulaceae

Crassu/a aquatica (L.) Schoenl. in Engl. & Prantl
Santa Catalina: RSA-POM.

Crassula erecta (H. & A.) Berger
San Miguel: SBBG; SBM.

Santa Rosa: Munz & Crow 1 1630 LA!; RSA-POM; SBM.
Santa Cruz: Fosberg 7621 LAM!; Fosberg 7643 LAM!;

RSA-POM; SBM.

Anacapa: SBBG.

San Nicolas: Newman 100 Pacif. Missile Test Center, Pt.
Mugu!; RSA-POM.

Santa Barbara: Dunkle 741 1 LAM!; RSA-POM; SBM.
Santa Catalina: Dunkle 2139 LAM!; Templeton 11395
LAM!; RSA-POM; SBM.

San Clemente: Dunkle 7323 LAM!; Moran 568 LAM!;

RSA-POM; SBM.

Guadalupe: Carlquist 445 RSA!.

Dudleya blochmanae (Eastw.) Moran ssp. insularis (Moran)
Moran

Santa Rosa: RSA-POM; SBBG; SBM.

Dudleya caespitosum (Haw.) Britt. & Rose
Santa Cruz: RSA-POM.

Anacapa: SBBG.

Dudleya candelabrum Rose

Santa Rosa: Moran 820 LAM!; SBBG; SBM.

Santa Cruz: Clokey 4949 LAM!; Hoffmann s.n. Apr. 10,
1930 SBM!; Balls & Blakley 23726 RSA!; CAS; SBBG.
Dudleya greenei Rose

San Miguel: Dunkle 8368 LAM!; Moran 3443 NY!; Hoff-
mann s.n. Jun. 11, 1930 SBM!; RSA-POM; SBBG.
Santa Rosa: Moran 810 LAM!; Moran 821 LAM!; Hoff-
mann s.n. Jun. 1 1, 1930 SBM!; RSA-POM; SBBG.
Santa Cruz: Clokey 5358 LAM!, LA!, NY!; Hoffmann s.n.
May 14, 1927 SBM#338!; Greene s.n. inJul.-Aug. 1886
CAS#244!; RSA-POM; SBBG.

Anacapa: Dunkle 7657 LAM!, AHFH!; Hoffmann s.n. Sep.

22, 1930 SBM#10912!.

Santa Catalina: RSA-POM; SBBG.

Dudleya guadalupensis Moran
Guadalupe: Moran 17428 MO!.

Dudleya hassei (Rose) Moran

Santa Catalina: Dunkle 2467 AHFH!; Trask s.n. in Feb.
1896 MO!; McClatchie s.n. 9/6/1893 NY!; RSA-POM;
SBBG; SBM.

Dudleya nesiotica (Moran) Moran

Santa Cruz; Moran 3362 CAS!; RSA-POM; SBBG; SBM.
Dudleya traskiae (Rose) Moran
Santa Barbara: Dunkle 8102 LAM!, AHFH!; RSA-POM;
SBBG.

Dudleya virens (Rose) Moran
San Nicolas: RSA-POM; SBBG.

Santa Catalina: Fosberg S4502 LAM!, NY!; Dunkle 2466
AHFH!; Fosberg S4890 LAM!; RSA-POM; SBBG.

San Clemente: Trask 165 NY!; RSA-POM; SBBG.
Guadalupe: Moran 6123 SD!; Moran 6131 SD!; Moran
13796 SD!.

Crossosomataceae
Crossosoma ca/ifornicum Nutt.

Santa Catalina: Trask s.n. in Jun. 1901 LAM!; Dunkle
2862 LAM!, LA!; Thome 39352 LAM!; RSA-POM;
SBBG; SBM.

San Clemente: RSA-POM.

Guadalupe: Moran 6630 RSA!; Moran 15194 RSA!.
Cucurbitaceae

Cucurbita foetidissima HBK.

Santa Cruz: RSA-POM; SBBG; SBM.

Marah guadalupensis (Wats.) Greene
Guadalupe: Anthony 234 LAM!; Palmer 33 CM!; Moran
5974 DS!.

Marah macrocarpus (Greene) Greene

San Miguel: Dunkle 8402 LAM!; RSA-POM; SBBG; SBM.
Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7622 LAM!, LA!; Fosberg 7584 LAM!;
Ellison s.n. May 12-15, 1929 LA!; RSA-POM; SBBG;
SBM.

Anacapa: SBBG; SBM.

San Nicolas: Dunkle 8359 LAM!; SBBG.

Santa Barbara: Dunkle 7439 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

Santa Catalina: RSA-POM; SBM.

San Clemente: Dunkle 7251 LAM!; Moran 573 LAM!;
Elmore 422 AHFH!.

Elatinaceae

Elatine californica Gray

Santa Catalina: Thome 35898 RSA!; SBBG.

Ericaceae

Arbutus menziesii Pursh

Santa Cruz: Hoffmann 599 LAM!; SBBG; SBM.
Arctostaphylos catalinae P.V. Wells

Santa Catalina: Wallace & Haefs 1412 RSA!; Trask s.n. in
Feb. 1900 ND-G#037709!; Fosberg S5398 LAM!, POM!;
SBBG; SBM.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 61

Arctostaphylos confertiflora Eastw.

Santa Rosa: Munz & Crow 1 1587 POM!, LA!; Moran 789
RSA!; Blakley 3168 RSA!; SBM.

Arctostaphylos insu/aris Greene f. insularis

Santa Cruz: Greene s.n. in Jul.-Aug. 1 886 ND-G#037684!,
ND-G#037685!, CAS#261!, CAS#261B!.
Arctostaphylos insularis Greene f. pubescens (Eastw.) P.V.
Wells

Santa Cruz: Fosberg7588 LAM!, LA!; Hutchinson s.n. Sep.
1, 1928 LAM!; Clokey 5022 LA!; RSA-POM; SBM.
Arctostaphylos tomentosa (Pursh) Lindl. ssp. insulicola P.V.
Wells

Santa Rosa: Hoffmann 101 Dec. 5, 1920 POM!.

Santa Cruz: Dunkle 8614 (in part) LAM!; Thome & Everett
36783 RSA!; Raven & Smith 1 529 1 RSA!; SBBG; SBM.
Arctostaphylos tomentosa (Pursh) Lindl. ssp. subcordata
(Eastw.) P.V. Wells

Santa Rosa: Moran 789 LAM!; Dunkle 8498 LAM!; RSA-
POM.

Santa Cruz: Dunkle 8614 (in part) LAM!; Howell 6335
CAS!; Greene s.n. in Jul.-Aug. 1886 ND-G#037697!;
RSA-POM; SBBG; SBM.

Arctostaphylos viridissima (Eastw.) McMinn

Santa Cruz: Howell 6368 CAS!; Balls & Blakley 23701
RSA!; Ellison s.n. May 12-15, 1929 LA!; SBBG.
Comarostaphylis diversifolia (Parry) Greene ssp. planifolia
(Jeps.) Wallace ex Thome

Santa Rosa: Thome et al. 4882 1 RSA!; Thome et al. 48989
RSA!; Munz & Crow 1 1662 POM!; SBBG; SBM.

Santa Cruz: Fosberg 7589 LAM!, LA!; Wolf 2827 RSA!;

Clokey 5019 LA!; ND-G; SBBG; SBM.

Santa Catalina: Thome & Everett 35032 RSA!, LA!; Dun-
kle 1712 POM!; Wallace 1404 RSA!; SBBG.

Vaccinium ovatum Pursh

Santa Rosa: Moran 8 1 3 LAM!; Epling & Erickson s.n. Aug.

8, 1937 LA!; RSA-POM; SBBG; SBM.

Santa Cruz: Fausett 6 LA!; RSA-POM; SBBG; SBM.

Xylococcus bicolor Nutt.

Santa Catalina: Trask s.n. in Feb. 1900 LAM!; Moran 650
LAM!; Thome 35969 RSA!; SBBG.

Euphorbiaceae

Eremocarpus setigerus (Hook.) Benth.

Santa Rosa: SBBG.

Santa Cruz: Hoffmann s.n. Aug. 7, 1930 POM!; Balls &
Blakley 23652 RSA!; Dunkle 8650 RSA!; SBBG.

Santa Catalina: Fosberg S4949 LAM!; Fosberg S4698
LAM!, POM!; Thome & Everett 34945 RSA!; SBBG;
SBM.

San Clemente: DeBuhr & Wallace 706 LAM!; House &
Grumbles s.n. Aug. 5-13, 1930 LAM!; Raven 17676
RSA!; SBBG.

Euphorbia crenulata Engelm.

Santa Catalina: Nuttall 170 F!; Nuttall 236 F!; Millspaugh
4867 F!.

Euphorbia melanadenia Torr.

Guadalupe: (Wheeler 1934: Palmer 783 at F).

Euphorbia misera Benth.

Santa Catalina: Trask s.n. in Oct. 1897 US!; Trask s.n. in
Dec. 1900 (scrap) NY!.

San Clemente: Dunkle 7336 LAM!; Blakley 632 RSA!;

Thome 36042 RSA!; SBBG.

Guadalupe: Rempel 759-37 LAM!; Moran 6136 RSA!.
Euphorbia peplis L.

Santa Rosa: Thome et al. 48883 RSA!.

Santa Cruz: Pierson 1 1046 RSA!; Daily 522 SCIR!.

Santa Catalina: Fosberg S4530 LAM!; Thome & Everett
33442 RSA!; Thome 36697 RSA!.

San Clemente: Dunkle 7216 LAM!.

Euphorbia pondii Millsp.

Guadalupe: Palmer 883 ND-G!; Moran 17353 RSA!;
Howell 8331 CAS!.

Euphorbia serpyllifolia Pers. var. serpyllifolia

Santa Catalina: Fosberg S4912 LAM!; Thome & Propst
37685 RSA!; Thome 36654 RSA!.

Euphorbia spathulata Lam.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Thome
35774 RSA!; Thome 39371 RSA!.

San Clemente: Raven 17675 RSA!; SBBG.

Ricinus communis L.

San Nicolas: Wier & Beauchamp s.n. Jul. 4, 1978 RSA!.
Santa Catalina: Thome & Everett 34875 RSA!; Thome
39353 RSA!; SBBG.

San Clemente: DeBuhr & Wallace 696 RSA!.

Fabaceae

Acacia decurrens Willd.

Santa Catalina: SBBG.

Acacia melanoxylon R. Br.

Santa Cruz: RSA-POM; SBBG.

Santa Catalina: Lister & Powell s.n. Apr. 1928 LAM!; RSA-
POM; SBBG.

Astragalus curtipes Gray

San Miguel: Dunkle 8404 LAM!; Munz & Crow 11828
POM!; SBBG; SBM.

Santa Rosa: Blakley 3127 RSA!; Moran 3337 RSA!; Munz
& Crow 11749 POM!; SBBG; SBM.

Astragalus didymocarpus H. & A. var. didymocarpus
San Miguel: SBBG; SBM.

Santa Rosa: Moran 799 LAM!, NY!; RSA!; Munz & Crow
1 1595 POM!, LA!; Raven, Blakley & Omduff 14913
RSA!; SBBG; SBM.

Santa Cruz: Hoffmann s.n. Sep. 11, 1931 LAM!; Munz &
Crow 1 1 890 POM!; Raven & Smith 15303 RSA!; SBBG;
SBM.

Anacapa: SBBG; SBM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Thome
36357 RSA!; Thome & Everett 34601 RSA!.

San Clemente: Trask 206 NY!, DS!.

Astragalus gambelianus Sheld.

Santa Cruz: Raven & Smith 15185 RSA!; SBBG; SBM.
Santa Catalina: Davidson, A. s.n. Jun. 26, 1891 LAM!;
Fosberg S4390 LAM!, POM!; Dunkle 2 1 52 LAM!; RSA-
POM; SBBG.

62 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Astragalus miguelensis Greene

San Miguel: Dunkle 8372 LAM!; Elmore 3 1 9 AHFH!; Voss
s.n. Sep. 1, 1930 POM!; CAS; SBBG; SBM.

Santa Rosa: Moran 794 LAM!, RSA!; Thorne et al. 48970
RSA!; Munz & Crow 11560 POM!, LA!; SBBG; SBM.
Santa Cruz: Clokey 4987 POM!; Raven & Smith 15313
RSA!; Webster et al. 97 RSA!; SBBG; SBM.

Anacapa: Dunkle 7606 LAM!, RSA!; Johnstone s.n. Jun.

25, 1932 LAM!; Moran 726 LAM!, RSA!; SBBG; SBM.
San Clemente: Beauchamp 323 LAM!, RSA!; DeBuhr &
Wallace 685 RSA!; Thorne 42904 RSA!; SBBG.
Astragalus nevinii Gray

San Clemente: Elmore 395 LAM!; Raven 18007 RSA!;
Thorne 35994 RSA!; SBBG.

Astragalus traskiae Eastw.

San Nicolas: Thome et al. 52358 RSA!; Raven & Thomp-
son 20712 RSA!; Trask s.n. in Apr. 1897 CAS!; LA;
SBBG.

Santa Barbara: Dunkle 8 1 27 LAM!, AHFH!; Dunkle 7436
LAM!; Dunkle 8132 AHFH!, RSA!; SBBG.

Astragalus trichopodus (Nutt.) Gray ssp. leucopsis (T. & G.)
Thome

Santa Rosa: Thorne et al. 48964 RSA!; SBM.

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 LA!; Williams
56 POM!; Raven & Smith 15292 RSA!; SBBG; SBM.
Anacapa: Ellison s.n. May 12-15, 1929 LA!; SBBG; SBM.
Santa Catalina: Dunkle 1 703 LAM!, POM!; Thorne & Ev-
erett 34979 RSA!; Jones s.n. May 1 5, 1903 POM!, MO!.
Astragalus trichopodus (Nutt.) Gray ssp. trichopodus

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Thorne &
Everett 34872 RSA!; Fosberg S443 1 LAM!; SBBG; SBM.
Coronilla valentina L.

Santa Catalina: SBBG.

Cytisus linifolius (L.) Lam.

Santa Catalina: Eaton s.n. Aug. 2, 1956 LAM!; Thome
36240 RSA!; Thome & Everett 33277 RSA!; SBBG.
Cytisus monspessu/anus L.

Santa Catalina: Thorne & Everett 34858 RSA!; Thorne
36309 RSA!; Henrickson 13814 RSA!; SBBG.

Lathyrus laetiflorus Greene ssp. alefeldii (White) Brads.
Santa Catalina: Fosberg S4806 LAM!, NY!, POM!; Trask
s.n. in Apr. 1897 LAM!; Thome 36205 RSA!; SBBG.
Lathyrus laetiflorus Greene ssp. barbarae (White) C.L. Hitchc.
Santa Rosa: Raven 1 4936 RSA!; Thome et al. 48993 RSA!;
SBBG; SBM.

Santa Cruz: Dunkle 8648 LAM!; Fosberg 7708 LAM!, NY!;

SBBG; SBM.

Anacapa: SBBG; SBM.

Santa Catalina: Fosberg S4378 POM!; Thome 35822 RSA!;

Knopf & Johnson 1454 RSA!; SBBG; SBM.

San Clemente: Trask 239 NY!; RSA-POM; SBBG.
Lathyrus tingitanus L.

Santa Catalina: Thome 36467 RSA!; Thome & Everett
34862 RSA!; Raven 17765 RSA!; SBBG.

Lathyrus vestitus Nutt, ex T. & G. ssp. vestitus
Santa Rosa: Munz & Crow 1 1578 POM!; Raven, Blakley
& Omduff 15012a RSA!.

Santa Cruz: Munz & Crow 1 1 836 POM!; Thorne & Everett
36827 RSA!; Raven & Smith 15217 RSA!.

Lotus argophyllus (Gray) Greene ssp. adsurgens (Dunkle)
Raven

San Clemente: Dunkle 7200 LAM!, RSA!; Elmore 410
AHFH!, LAM!.

Lotus argophyllus (Gray) Greene ssp. niveus (Greene) Munz

Santa Cruz: Abrams & Wiggins 199 RSA!; Greene s.n. in
Jul.-Aug. 1886 CAS#680!; Hoffmann s.n. Sep. 22, 1930
POM!; SBBG; SBM.

Lotus argophyllus (Gray) Greene ssp. ornithopus (Greene)
Raven

San Nicolas: Trask 42 LAM!; Wallace et al. 1609 LAM!;
Foreman, Evans & Rainey 71 LA!; CAS; RSA-POM;
SBBG; SBM.

Santa Barbara: Dunkle 7409 LAM!; Moran 825 AHFH!;
Kanakoff s.n. in Aug. 1940 LAM!; RSA-POM; SBBG;
SBM.

Santa Catalina: Grant 7162 LAM!, POM!, USC!; Thorne
34900 RSA!; Fosberg 81 1 1 LA!; SBBG; SBM.

San Clemente: Munz 6611 LAM!, POM!; Dunkle 7350
AHFH!; Raven 17149 RSA!; SBBG.

Guadalupe: Howell 8177 MO!, POM!; Moran 5951 LA!,
RSA!; Greene s.n. Apr. 19, 1885 CAS#681!.

Lotus corniculatus L.

Santa Cruz: Daily 150 SCIR!.

Santa Catalina: Thorne 36700 RSA!; SBBG.

Lotus grandiflorus (Benth.) Greene var. grandiflorus

Santa Rosa: Munz & Crow 1 1638 POM!; SBM.

Santa Cmz: Thome & Everett 36751 RSA!; Breedlove 2824
RSA!; SBBG; SBM.

Santa Catalina: Moran 627 LAM!, RSA!; Dunkle 1749
AHFH!, POM; Thorne 35912 RSA!; SBBG.

Guadalupe: Palmer 23 MO!; Greene s.n. in Apr. 1 885 ND-
G#025788!; Greene s.n. in Apr. 1885 ND-G#025789!.

Lotus hamatus Greene

Santa Rosa: Hoffmann 4 1 8 POM!; Hoffmann s.n. Mar. 23,
1929 POM!; SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Davidson s.n. Jun. 26, 1 89 1 LAM!; Dunkle
1841 AHFH!, POM!; Thorne 35843 RSA!; LA; MO;
SBBG; SBM.

San Clemente: Raven 17591 RSA!; SBBG.

Lotus heermannii (Dur. & Hilg.) Greene ssp. heermannii

Santa Catalina: (Ottley 1923: G.B. Grant 716 no herbar-
ium cited).

Lotus humistratus Greene

Santa Cruz: Pierson, F.W. 1 1084 RSA!; SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Trask s.n.
in Apr. 1898 US!.

Lotus micranthus Benth.

Santa Cruz: Pierson 7133 RSA!; Wolf 2854 RSA!; SBM.

Lotus purshianus (Benth.) Clem. & Clem. ssp. purshianus

Santa Cruz: Weissman 325 DS!; Gorelick s.n. Jun. 28, 1978
RSA!.

Santa Catalina: Fosberg S4386 LAM!, POM!; Thome &
Everett 34074 RSA!; Dunkle 2432 AHFH!; SBBG; SBM.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 63

Lotus salsuginosus Greene ssp. salsuginosus

San Miguel: SBBG; SBM.

Santa Rosa: Munz & Crow 1 1 758 POM!, LA!; SBBG; SBM.

Santa Cruz: Fosberg 7647 LAM!; Fosberg 7618 LAM!,
LA!; Blakley 3384 RSA!; SBBG; SBM.

Anacapa: SBBG.

Santa Catalina: Fosberg S4415 LAM!, POM!; Fosberg
S4422 LAM!; Trask s.n. in Mar. 1896 MO# 1864670!;
RSA-POM; SBBG.

Lotus scoparius (Nutt, in T. & G.) Ottley var. dendroideus
(Greene) Ottley

San Miguel: SBBG.

Santa Rosa: Dunkle 8440 LAM!, RSA!; Elmore 196
AHFH!; Munz & Crow 1 1 599 LA!, POM!; SBBG; SBM.

Santa Cruz: Moran 768 LAM!, RSA!; Clokey 5191 LA!,
RSA!; Greene s.n. in Jul.-Aug. 1886 CAS#673!,
DS#35523!; SBBG; SBM.

Anacapa: Dunkle 76 1 7 LAM!, AHFH!; Gustafson s.n. Apr.
9, 1973 LAM!; Moran 733 LAM!, RSA!; SBBG; SBM.

Santa Catalina: Fosberg S4940 LAM!; Fosberg 8115 LA!,
POM!; Heller 8952 NY!; SBBG; SBM.

San Clemente: Wiggins 1 1956 RSA!; SBBG.

Lotus scoparius (Nutt, in T. & G.) Ottley ssp. scoparius

San Miguel: SBBG; SBM.

Santa Rosa: SBBG; SBM.

Santa Cruz: Greene s.n. in Jul.-Aug. 1 886 ND-G#025862!.

Anacapa: SBBG.

Santa Catalina: Dunkle 1851 AHFH!; SBBG.

Lotus scoparius (Nutt, in T. & G.) Ottley ssp. traskiae (Eastw.
ex Noddin in Abrams) Raven

San Clemente: Dunkle 7285 LAM!, RSA!; Elmore s.n. Nov.
26, 1939 AHFH!; Dunkle 7281 LAM!; SBBG.

Lotus scoparius (Nutt, in T. & G.) Ottley var. veatchii (Greene)
Ottley

San Miguel: Munz & Crow 1 1780 POM!; Moran 3442
POM!; Greene s.n. in Sep. 1886 CAS#682!; SBM.

Santa Cruz: Greene s.n. in Jul.-Aug. 1886 ND-G#025868!.

Lotus strigosus (Nutt, in T. & G.) Greene ssp. strigosus

San Miguel: SBM.

Santa Rosa: Munz & Crow 1 1633 POM!; Raven, Blakley
& Omduff 14960 RSA!; SBBG; SBM.

Santa Cruz: Wolf 2748 RSA!; Thome & Everett 36838
RSA!; Raven & Smith 15273 RSA!; SBBG; SBM.

Anacapa: SBBG; SBM.

Santa Catalina: Fosberg S4393 LAM!, POM!; Moran 701
LAM!; Thome 35871 RSA!; SBBG.

San Clemente: Munz 6750 POM!; Raven 17705 RSA!;
Thome 42798 RSA!; SBBG.

Lotus subpinnatus Lag.

Santa Rosa: Raven 14932 RSA!; SBM.

Santa Cruz: Fosberg 7644 LAM!; Hoffmann s.n. Apr. 12,
1 93 1 LAM!; Munz & Crow 11518 POM!, LA!; SBM.

Anacapa: SBM.

Santa Catalina: Trask s.n. in Apr. 1900 LAM!; Fosberg
S4465 LAM!; Dunkle 2100 AHFH!; RSA-POM.

Lupinus agardhianus Heller

Santa Rosa: Thome et al. 48978 RSA!; Munz & Crow
11634 POM!; Hoffmann Apr. 17, 1921 POM!; SBM.

Santa Cruz: Elmore s.n. Apr. 19, 1936 LAM!, AHFH!;
Munz & Crow 11511 POM!; Clokey 4976 US!; SBBG;
SBM.

Santa Catalina: Fosberg S4413 LAM!, US!; Trask s.n. in
May 1897 US!; Dunkle 1835 AHFH!, POM!; LA; SBM.
San Clemente: Thome 42797 RSA!; Thome 42947 RSA!;
Raven 17706 RSA!; SBBG.

Lupinus albifrons Benth. var. albifrons
San Miguel: Dunkle 8387 LAM!; RSA!; SBBG; SBM.
Santa Rosa: Moran 805 LAM!, RSA!; Munz & Crow 11752
LA!, POM!; Raven, Blakley & Omduff 14958 RSA!;
SBBG; SBM.

Santa Cruz: Balls & Blakley 23689 RSA!; Thome & Everett
36826 RSA!; Webster et al. 73 RSA!; SBBG.

Anacapa: Dunkle 7621 AHFH!; Moran 738 LAM!, RSA!;

Hoffmann s.n. Mar. 16, 1929 POM!; SBBG; SBM.

San Nicolas: Trask s.n. in Mar. 1901 LAM!; Kanakoff s.n.
Apr. 14, 1940 LAM!; Foreman & Smith 186 LA!; RSA-
POM; SBBG; SBM; US.

Santa Catalina: Moran 704 LAM!, RSA!; Eastwood 6472
US!; Thome 33439 RSA!; LA; MO; SBBG.

Lupinus albifrons Benth. var. douglasii (J.G. Agardh) C.P.
Sm.

San Miguel: SBBG.

Santa Rosa: Thome et al. 48969 RSA!; Thome et al. 48904
RSA!; Thome et al. 49033 RSA!; SBM.

Santa Cruz: Hoffmann s.n. Mar. 22, 1929 SBM!; Wolf
2770 RSA!; Raven & Smith 15281 RSA!; SBBG; SBM.
San Nicolas: Foreman, Evans & Rainey 28 RSA!; Raven
& Thompson 20759 RSA!; Raven & Thompson 20723
RSA!; SBBG.

Lupinus arboreus Sims

San Miguel: Dunkle 8383 LAM!, AHFH!, MO!, RSA!;
Blakley 5854 SBBG!; SBM.

Santa Rosa: Moran 796 LAM!, MO!, RSA!; Munz & Crow
11760 LA!, POM!; SBBG.

Lupinus bicolor Lindl. ssp. microphyllus (Wats.) D. Dunn
San Miguel: SBBG.

Santa Rosa: Hoffmann s.n. Apr. 16, 1929 POM!; Dunn,
N. s.n. May 15, 1932 LA!; Raven, Blakley & Omduff
14901 RSA!; SBBG.

Santa Cruz: Elmore 446 LAM!, AHFH!; Webster et al. 87
RSA!; Raven & Smith 15137 RSA!; SBBG.

Anacapa: SBBG.

San Nicolas: Trask s.n. in Apr. 1897 US!.

Santa Catalina: Fosberg S4377 LAM!, POM!, US!; Nuttall
169 F!; Dunkle 1801 AHFH!; SBBG.

San Clemente: Moran 592 LAM!, RSA!; Dunkle 7321
AHFH!; SBBG.

Guadalupe: Moran 5959 RSA!; Norris s.n. May 1, 1951
LA!.

Lupinus bicolor Lindl. ssp. tridentatus (Eastw. ex C.P. Sm.)
D. Dunn

Santa Cruz: Wolf 2747 RSA!.

Lupinus bicolor Lindl. ssp. umbellatus (Greene) D. Dunn
San Miguel: Munz & Norris 1 1775 POM!.

Santa Rosa: Munz & Crow 1 1592 POM!.

64 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Santa Cruz: Fosberg 7530 LAM!, LA!, POM!; Clokey 4977
NY!, POM!; Fosberg 7657 LAM!; SBBG.

Santa Catalina: Lister & Powell s.n. Apr. 2, 1928 LAM!;
Dunkle 1741 POM!.

San Clemente: Dunkle 7319 LAM!; Munz 6742 POM!;
Trask 256 US!; SBBG.

Lupinus concinnus J.G. Agardh ssp. concinnus
Santa Cruz: Wolf 2802 RSA!; Elmore s.n. Apr. 19, 1936
LAM!; Pierson 1 1065 RSA!.

Santa Catalina: Merritt s.n. in Aug. 1894 LAM!; Thorne
& Everett 34469 RSA!; Thorne 35946 RSA!; SBBG.

Lupinus densiflorus Benth. var. palustris (Kell.) C.P. Sm.
Santa Rosa: Hoffmann 729 POM!; Blakley 3189 RSA!;
Thorne et al. 48907 RSA!.

Santa Cruz: Sauer et al. 5494 RSA!; Raven & Smith 15311
RSA!; Webster et al. 101 RSA!.

Lupinus guadalupensis Greene

San Clemente: Moran 587 LAM!, MO!, NY!; Beauchamp
262 RSA!; Munz 6741 POM!; SBBG.

Guadalupe: Carlquist 447 RSA!; Greene s.n. Apr. 23, 1885
CAS#702!.

Lupinus hirsutissimus Benth.

Santa Rosa: SBM.

Santa Cruz: Wolf 2727 RSA!; Munz & Crow 1 1 538 POM!;

Thome & Everett 36835 RSA!; SBBG; SBM.

Santa Catalina: Trask s.n. in May 1 898 LAM!; Moran 694
LAM!; Dunkle 1936 AHFH!; RSA-POM.

San Clemente: Dunkle 7299 LAM!, AHFH!, RSA!; Trask
255 US!; Raven 17702 RSA!; SBBG.

Lupinus niveus Wats.

Guadalupe: Moran 17348 LAM!; Moran 6475 DS!;
Carlquist 474 RSA!.

Lupinus polycarpus Greene
Santa Rosa: SBM.

Lupinus succulentus Dougl. ex Koch

San Miguel: Munz & Voss 11797 POM!; SBBG; SBM.
Santa Rosa: Moran 803 LAM!, RSA!; Thome et al. 48965
RSA!; Munz & Crow 11716 POM!; SBBG; SBM.

Santa Cruz: Fosberg 7537 LAM!, LA!; Wolf 2812 RSA!;

Webster et al. 70 RSA!; SBBG; SBM.

Anacapa: Moran 712 LAM!, RSA!.

Santa Catalina: Fosberg S4414 LAM!; Moran 676 LAM!;

Trask s.n. in Apr. 1 895 US!; AHFH; RSA-POM; SBBG.
San Clemente: Dunkle 7279 LAM!; Thorne 42906 RSA!;
Raven 17677 RSA!; SBBG.

Lupinus truncatus Nutt, ex H. & A.

Santa Rosa: Thome et al. 48731 RSA!; Raven, Blakley &
Omduff 14973 RSA!; SBBG; SBM.

Santa Cruz: Hoffmann s.n. Jun. 15, 1930 LAM!; Clokey
4975 POM!, US!; Fosberg 7673 LAM!; SBBG; SBM.
Anacapa: Moran 728 LAM!, SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Trask s.n.

in May 1897 US!; Moran 692 LAM!, RSA!; SBBG.

San Clemente: Dunkle 7303 LAM!; Moran 588 LAM!,
RSA!; Trask 257 US!; SBBG.

Medicago polymorpha L. var. brevispina (Benth.) Heyn.
Santa Rosa: SBBG.

Santa Cruz: Dunkle 8573 LAM!; Hoffmann s.n. Jun. 15,
1930 LAM!; Raven & Smith 15144 RSA!; SBBG.
Santa Catalina: Dunkle 1 8 1 4 AHFH!; Thorne 3690 1 RSA!;

Lister & Powell s.n. Apr. 5, 1928 USC!.

San Clemente: Raven 17131 RSA!.

Guadalupe: Moran 6763 SD!; Moran 6783 RSA!; Wiggins
& Ernst 94 DS!.

Medicago polymorpha L. var. polymorpha

San Miguel: Munz & Crow 1 1872 POM!; SBBG.

Santa Rosa: Dunkle 8437 LAM!; Raven, Blakley & Orn-
duff 14888 RSA!; Thorne et al. 49047 RSA!; SBBG;
SBM.

Santa Cruz: Thorne & Everett 36848 RSA!; Balls & Blakley
23737 RSA!; Raven & Smith 15143 RSA!; SBBG; SBM.
Anacapa: SBBG.

San Nicolas: Dunkle 8323 LAM!; Raven & Thompson
20709 RSA!; Thorne et al. 52365 RSA!; SBBG.

Santa Barbara: Dunkle 7463 LAM!, AHFH!, RSA!; Thorne
37498 RSA!; SBBG.

Santa Catalina: Fosberg S432 1 LAM!; Lister & Powell s.n.

USC!; Thorne 35929a RSA!; SBBG.

San Clemente: Dunkle 7327 LAM!; Munz 6691 POM!;
Raven 17130 RSA!.

Guadalupe: Moran 2884 DS!; Wiggins & Ernst 93 DS!.
Medicago saliva L.

Santa Rosa: SBBG.

Santa Cruz: Dunkle 8572 LAM!, RSA!; Hoffmann s.n. Jun.

15, 1930 LAM!; SBBG; SBM.

Anacapa: Dunkle 7465 LAM!; SBBG.

San Nicolas: Trask s.n. in Apr. 1901 LAM!.

Santa Catalina: Fosberg S4421 LAM!; Dunkle 1863
AHFH!, POM!; Thome & Everett 34635 RSA!.

San Clemente: Thome 42977 RSA!; SBBG.

Melilotus alba Medicus
Santa Cruz: SBBG.

San Nicolas: Foreman, Evans & Rainey 103 LA!; Thome
et al. 52368 RSA!; Raven & Thompson 20738 RSA!;
SBBG.

Santa Catalina: Fosberg S4603 LAM!; Thome 36268 RSA!;
SBBG.

San Clemente: Raven 17959 RSA!; SBBG.

Melilotus indica (L.) All.

San Miguel: Dunkle 8391 LAM!, RSA!; SBBG; SBM.
Santa Rosa: Thome et al. 48810a RSA!; Thorne et al.
49046 RSA!; SBM.

Santa Cruz: Dunkle 8571 LAM!; Williams 44 POM!; SBBG;
SBM.

Anacapa: SBBG; SBM.

San Nicolas: Dunkle 8337 LAM!, RSA!; Kanakoff s.n. Apr.
18, 1940 LAM!; Foreman, Evans & Rainey 88 LA!;
SBBG.

Santa Catalina: Fosberg S4322 LAM!; Dunkle 1985
AHFH!; Thorne 35764 RSA!; SBBG.

San Clemente: Raven 17960 RSA!; Raven 17128 RSA!;

Blakley 6368 RSA!; SBBG.

Guadalupe: Wiggins & Ernst s.n. DS!.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 65

Pickeringia montana Nutt. ssp. montana
Santa Cruz: Hoffmann s.n. May 23, 1932 POM!; SBBG.
Spartium junceum L.

San Nicolas: Newman 109 Pacific Missile Test Center, Pt.

Mugu!; Wier & Beauchamp s.n. Jun. 29, 1978 RSA!.
Santa Catalina: Thorne 36907 RSA!; SBBG.

Trifolium albopurpureum T. & G.

Santa Rosa: SBBG; SBM.

Santa Cruz: Hoffmann 2 1 3 POM!; Thome & Everett 36810
RSA!; Raven & Smith 15184 RSA!; SBBG; SBM.

San Nicolas: Trask 37 LAM!, NY!; Thome et al. 52337
RSA!.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Kennedy
10028 POM!; Thome & Everett 34603 RSA!.

Trifolium amplectens T. & G. var. amplectens
San Miguel: SBM.

Santa Rosa: Raven, Blakley & Omduff 14896 RSA!; SBBG;
SBM.

Santa Cruz: Hoffmann s.n. Sep. 10, 1931 LAM!; Thome
& Everett 36852 RSA!; Raven & Smith 15192 RSA!;
SBBG; SBM.

Santa Catalina: Dunkle 2 1 29 AHFH!; Dunkle 1 770 AHFH!;
Thome 35986 RSA!; SBBG.

San Clemente: Dunkle 7315 AHFH!; Thome 36 1 26 RSA!;
Raven 17613 RSA!; SBBG.

Guadalupe: Howell 8285 CAS!; Wiggins & Ernst 92 DS!.
Trifolium amplectens T. & G. var. truncatum (Greene) Jeps.
San Miguel: Munz & Crow 1 1830 POM!.

Santa Rosa: Munz & Crow 11711 POM!; Hoffmann s.n.
May 10, 1932 POM!; SBM.

Santa Cruz: Fosberg 7528 LAM!, LA!; Munz & Crow 1 1 520
POM!; Pierson 11030 POM!; SBM.

Anacapa: SBBG.

Santa Catalina: Dunkle 1 745 AHFH!, POM!; Kennedy s.n.
Apr. 21, 1907 LA!, POM!; Dunkle 1770 POM!; SBM.
Trifolium barbigerum Torr.

San Miguel: Munz & Norris 1 1764 POM!.

Santa Rosa: Hoffmann 711 POM!; Raven 14931 RSA!;
SBBG.

Trifolium ciliolatum Benth.

Santa Rosa: SBM.

Santa Cruz: Munz & Crow 1 1 557 POM!; Thome & Everett
36837 RSA!; Raven & Smith 15188 RSA!; SBBG.
Santa Catalina: Trask s.n. in May 1897 LAM!.

Trifolium depauperatum Desv.

Santa Cruz: Raven & Smith 13239 RSA!.

San Nicolas: Howell 8212 CAS!.

Trifolium fucatum Lindl. var. gambelii (Nutt.) Jeps.

San Miguel: Hoffmann 584 POM!; Hoffmann s.n.

SBM#1 1954!; SBBG.

Santa Rosa: Hoffmann 721 POM!; SBM.

Santa Cruz: SBBG; SBM.

Santa Catalina: Gambel s.n. GH!.

San Clemente: Raven 17674 RSA!; SBBG.

Trifolium gracilentum T. & G.

San Miguel: Munz & Norris 1 1767 POM!; SBM.

Santa Rosa: Munz & Crow 1 1702 POM!; SBBG; SBM.

Santa Cruz: Fosberg 7679 LAM!; Munz & Crow 11895
POM!; Raven & Smith 15134 RSA!; SBBG; SBM.
Anacapa: SBBG; SBM.

San Nicolas: Trask 35 MO!.

Santa Catalina: Dunkle 2126 AHFH!; Kennedy s.n. Apr.

21, 1907 LA!; Thome 34604 RSA!; SBBG.

San Clemente: Raven 17200 RSA!; Munz 6692 POM!;

Thome 42793b RSA!; SBBG.

Guadalupe: Howell 8247 POM!; Moran 6599 RSA!; Lind-
say s.n. Apr. 12, 1948 RSA!.

Trifolium macraei H. & A.

Santa Rosa: Munz & Crow 1 1 598 POM!, LA!; Raven 14945
RSA!; Hoffmann 686 POM!; SBM.

Santa Cruz: Wolf 2850 RSA!; Hoffmann 212 POM!; Munz
& Crow 11510 POM!; SBM.

Santa Catalina: Dunkle 2 1 08 AHFH!; Dunkle 1 839 AHFH!,
POM!; Fritchey, J.Q.A. 19 MO!.

Trifolium microcephalum Pursh
San Miguel: SBBG; SBM.

Santa Rosa: Blakley 3059 RSA!; Munz & Crow 11632
POM!; Raven, Blakley & Omduff 14898 RSA!; SBBG;
SBM.

Santa Cruz: Fosberg 7648 LAM!, LA!; Clokey 4979 NY!;

Elmore 441 AHFH!; RSA-POM; SBBG; SBM.

Santa Catalina: Fosberg S4342 LAM!, POM!; Trask s.n.

in Mar. 1900 NY!; Thome 36226 RSA!; SBBG; SBM.
San Clemente: Trask 336 NY!; Munz 6603 POM!; Pierson
3435 RSA!; SBBG.

Guadalupe: Palmer 27 NY!; Howell 8246 NY!; Rose 16036
NY!; RSA-POM.

Trifolium microdon H. & A. var. pilosum Eastw.

San Nicolas: Trask s.n. in Apr. 1897 CAS#1179!, US!;
Trask 34 MO!.

Santa Catalina: Trask s.n. in May 1897 MO!, US!; Trask
s.n. in May 1903 US!.

Trifolium palmeri Wats.

San Nicolas: Trask s.n. in Apr. 1897 US!; Trask 37 MO!.
Santa Barbara: Dunkle 7449 LAM!, AHFH!, RSA!; Dunkle
7415 LAM!; SBBG.

Santa Catalina: Trask s.n. in Mar. 1896 MO!; Trask 334
NY!; Thome 42793a NY!, DS!; SBBG.

San Clemente: Raven 17648 RSA!; Thome 42899 MO!;

Wooton s.n. Apr. 26, 1912 US!; DS; SBBG.
Guadalupe: Palmer 26 NY!, CM!; Howell 8308 CAS!, NY!;
Franceschi 27 RSA!.

Trifolium repens L.

Santa Catalina: Thome 36699 RSA!.

Trifolium tridentatum Lindl. var. aciculare (Nutt.) McDer.
San Miguel: SBM.

Santa Rosa: Munz & Crow 1 1596 LA!; Moran 798 RSA!;

Thome et al. 48747 RSA!; SBM.

Santa Cruz: Fosberg 7651 LAM!; Hoffmann s.n. LA!; El-
more 465 AHFH!; SBM.

Anacapa: SBBG; SBM.

San Nicolas: Raven & Thompson 20783 RSA!.

Santa Barbara: Dunkle 7442 LAM!, AHFH!; Dunkle 8115
AHFH!; Dunkle 7425 RSA!; SBBG.

66 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Santa Catalina: Dunkle 2112 AHFH!; Trask s.n. in Mar.

1901 LAM!; Thorne & Everett 34485 RSA!.

San Clemente: Dunkle 7288 LAM!, AHFH!, RSA!; Raven
17154 RSA!; Raven 17647 RSA!; SBBG.

Trifolium tridentatum Lindl. var. tridentatum
San Miguel: SBBG.

Santa Rosa: Moran 798 LAM!; Munz & Crow 1 1 596 POM!;

Munz & Crow 1 1672 POM!; SBBG.

Santa Cruz: Munz & Crow 1 1559 POM!; Blakley 3313
RSA!; Raven & Smith 15154 RSA!; SBBG; SBM.
Anacapa: SBBG.

San Nicolas: RSA-POM; SBBG.

Santa Barbara: Dunkle 7425 AHFH!, MO!; Thorne 37500
RSA!; Blakley 5678 RSA!; SBBG; SBM.

Santa Catalina: Fosberg S4367 (in part) LAM!; Kennedy
s.n. Apr. 21, 1907 LA!, RSA!; Thorne 36867 RSA!;
SBBG.

San Clemente: Thorne 36060 RSA!; Thorne 42870 RSA!;
SBBG.

Trifolium variegatum Nutt, in T. & G.

Santa Cruz: Hoffmann s.n. Jun. 15, I 930 POM#l 7201 1 !;
SBM.

Vicia americana Muhl. ex Willd.

San Miguel: SBBG; SBM.

Santa Rosa: Munz & Crow 1 1677 POM!, LA!; Thorne et
al. 48765 RSA!; Hoffmann s.n. Apr. 16, 1929
SBM#6195!; SBBG.

Santa Cruz: Raven & Smith 15157 RSA!; Pierson, F.W.
1 1061 RSA!; SBBG; SBM.

Vicia dasycarpa Ten.

San Nicolas: Foreman, Evans & Rainey 93 LA!; SBBG.
Santa Catalina: Parratt 546 LAM!; Thorne 36498 RSA!;
SBBG.

Vicia exigua Nutt, in T. & G.

Santa Rosa: SBM.

Santa Cruz: Brandegee s.n. in 1888 (in part) UC!; Hoff-
mann s.n. Sep. 21,1 930 POM!; Thome & Everett 36843
RSA!; SBBG.

Anacapa: SBM.

Santa Catalina: Fosberg 4617 LAM!, POM!, UC!; Dunkle
1 767 AHFH!, POM!; Thome 35954 RSA!; SBBG; SBM.
San Clemente: Raven 17338 RSA!; Raven 17680 RSA!,
UC!; SBBG.

Guadalupe: Moran 1 7478 RSA!, UC!; Newcomb 1 88 UC!.
Vicia hassei Wats.

Santa Rosa: Moran 782 LAM!, RSA!; Munz & Crow 11614
POM!; SBBG.

Santa Cruz: Munz & Crow 1 1845 POM!; Pierson 1 1085
RSA!; Abrams & Wiggins 8 UC!.

Anacapa: [Hoffmann] s.n. Mar. 1 1, 1928 SBM#4158!.

San Nicolas: Trask 85 MO!, NY!.

San Clemente: Munz 6669 POM!, UC!; Pierson 3442 RSA!;

Thome 42962 RSA!.

Guadalupe: Wiggins & Ernst 200 UC!.

Vicia villosa Roth

San Nicolas: Raven & Thompson 20766 RSA!; SBBG.

Fagaceae

Quercus agrifolia Nee var. agrifolia
Santa Rosa: Dunkle 8494 LAM!, AHFH!; Thorne et al.
48771 RSA!; Epling & Erickson s.n. Aug. 8, 1937 LA!;
SBM.

Santa Cruz: Fosberg 7628 LAM!; Clokey 4895 LAM!,
POM!; Raven & Smith 15205 RSA!; SBM.

Quercus chrysolepis Liebm.

Santa Cruz: Clokey 4897 LAM!; Clokey 4900 LAM!; Hoff-
mann s.n. Jun. 15, 1930 LAM!; SBBG; SBM.

Santa Catalina: Trask s.n. in Apr. 1900 LAM!; Thorne &
Thome 36420 RSA!; Thorne & Thorne 36430 RSA!.
San Clemente: Dunkle 7361 LAM!, AHFH!, RSA!.
Quercus douglasii H. & A.

Santa Cruz: Thome & Everett 36774 RSA!; Raven & Smith
15277 RSA!.

Santa Catalina: SBBG.

Quercus dumosa Nutt.

Santa Rosa: Dunkle 8487 LAM!, RSA!; Thome et al. 48855
RSA!; Munz & Crow 1 1577 POM!; SBM.

Santa Cruz: Clokey 4896 LAM!, LA!, POM!; Dunkle 8659
LAM!, RSA!; Raven & Smith 15160 RSA!; SBM.

Santa Catalina: Fosberg S4358 LAM!; Grant 6137 LAM!;
Moran 707 LAM!, RSA!; SBBG; SBM.

Quercus engelmannii Greene
Santa Catalina: Thome & Everett 34611 RSA!; Thome
36864 RSA!.

Quercus lobata Nee

Santa Cruz: Thome & Everett 36776 RSA!; SBBG.

Santa Catalina: SBBG.

Quercus x macdonaldii Greene
Santa Rosa: SBM.

Santa Cruz: Fosberg 7558 LAM!; Hoffmann s.n. Jun. 15,
1930 LAM!; Clokey 5173 LA!, RSA!; SBM.

Santa Catalina: Fosberg S4589 LAM!; Wolf 4230 RSA!;
Thome 36189 RSA!; SBM.

Quercus x morehus Kell.

Santa Cruz: Wolf 2759 RSA!; Pierson 1 1072 RSA!; SBM.
Quercus tomentel/a Engelm.

Santa Rosa: Dunkle 8507 LAM!; Epling & Erickson s.n.

Apr. 8, 1 938 LA!; Moran 8 1 5 LAM!, RSA!; SBBG; SBM.
Santa Cruz: Fosberg 7594 LAM!; Munz & Crow 11501
LA!, POM!; Raven & Smith 15159 RSA!; SBBG; SBM.
Anacapa: Dunkle 7660 LAM!, AHFH!, RSA!; Moran 714
LAM!, RSA!; SBBG; SBM.

Santa Catalina: Fosberg S5607 LAM!; Raven 17808 LA!;
Thome 35823 RSA!; SBBG.

San Clemente: DeBuhr & Wallace 700 LAM!, RSA!; Dun-
kle 7357 LAM!, AHFH!; Raven 17720 RSA!; SBBG.
Guadalupe: Moran 18387 LAM!; Moran 13788 LAM!;
Walker s.n. Nov. 12, 1938 AHFH!; RSA-POM.
Quercus wis/izenii A. DC. var . frutescens Engelm.

Santa Cruz: Sauer & Mathias 5676 RSA!; SBM.
Frankeniaceae

Frankenia grandifolia Cham. & Schlecht. var. grandifolia
San Miguel: Dunkle 8411 LAM!, RSA!; Dunkle 8382
AHFH!; SBBG; SBM.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 67

Santa Rosa: Dunkle 8430 LAM!; Blakley 3 1 96 RSA!; Munz
& Crow 11564 OM!; SBBG; SBM.

Santa Cruz: Clokey 5005 POM!; SBBG; SBM.

Anacapa: Dunkle 7603 LAM!, AHFH!, RSA!; Elmore 250
AHFH!; Dunn, N. s.n. May 14, 1932 LA!; SBBG; SBM.

San Nicolas: Dunkle 8300 LAM!; Kanakoff s.n. Apr. 22,
1 940 LAM!; Raven & Thompson 20690 LA!; RSA-POM;
SBBG.

Santa Catalina: Fosberg S4471 LAM!; Fosberg S5424
LAM!; Dunkle 1923 AHFH!; RSA-POM; SBBG; SBM.

San Clemente: House & Grumbles s.n. Aug. 5-13, 1930
USC!; Thome 42912 RSA!; Raven 17183 RSA!; SBBG.

Guadalupe: Moran 6472 RSA!.

Garryaceae

Garrya elliptica Dougl.

Santa Cruz: Williams 55 POM!; Bartholomew & Zadnik
712b RSA!; Beeks 28-4-68-17 RSA!; SBBG.

Gentianaceae

Centaurium davyi (Jeps.) Abrams

Santa Rosa: Elmore 204 AHFH!.

Santa Cruz: Blakley 3296 RSA!; Hoffmann s.n. Jun. 10,
1930 POM!; Clokey 5117 NY!.

San Clemente: Trask 174 US!.

Centaurium exaltatum (Griseb.) W. Wight

Santa Cruz: Greene s.n. in Jul.-Aug. 1 886 ND-G#03892 1 !,
US#3 10252!.

Centaurium muhlenbergii (Griseb.) W. Wight

Santa Cruz: (Jepson 1909-1943: Jepson 12089).

Centaurium venustum (Gray) Rob.

Santa Catalina: Dunkle 1905 AHFH!, POM!; Thome &
Everett 34914 RSA!; Thome 36354a RSA!; SBBG.

Geraniaceae

Erodium botrys (Cav.) Bertol.

Santa Rosa: Munz & Crow 1 1636 POM!; SBM.

Santa Cruz: Thome & Everett 3683 1 RSA!; Raven & Smith
15172 RSA!.

San Clemente: Thome 36083 RSA!.

Erodium cicutarium (L.) L’Her.

San Miguel: Dunkle 8378 LAM!, AHFH!; Munz & Crow
11804 POM!; SBBG; SBM.

Santa Rosa: Thome et al. 48756 RSA!; Dunkle 8464 LAM!;
SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Anacapa: SBBG; SBM.

San Nicolas: Dunkle 8338 LAM!, AHFH!; Foreman 228
LA!; Thome et al. 52370 RSA!; SBBG; SBM.

Santa Barbara: Dunkle 7462 LAM!, AHFH!, RSA!; Dunkle
8134 LAM!; Dunkle 8120 AHFH!; RSA-POM; SBBG.

Santa Catalina: Fosberg S4532 LAM!; Fosberg S4318
LAM!; Thome & Everett 33459 RSA!.

San Clemente: House & Grumbles s.n. Aug. 5-13, 1930
USC!; Munz 6704 POM!; Raven 17136 RSA!; SBBG.

Guadalupe: Palmer 19 CM!; Wiggins & Ernst 60 DS!.

Erodium macrophyllum H. & A. var. californicum (Greene)
Jeps.

Santa Cmz: Brandegee s.n. in Apr. 1888 UC!.

Erodium moschatum (L.) L’Her.

San Miguel: SBBG; SBM.

Santa Rosa: Thome et al. 48738 RSA!; Thome et al. 49015
RSA!; Raven, Blakley & Omduff 14881 RSA!; SBBG;
SBM.

Santa Cmz: Dunkle 8627 LAM!; Thome & Everett 36845
RSA!; Webster et al. 88 RSA!; SBBG; SBM.

Anacapa: SBBG.

San Nicolas: Kanakoff s.n. Apr. 14, 1940 LAM!; Thome
et al. 52369 RSA!; Raven & Thompson 20781 RSA!;
SBBG.

Santa Barbara: Dunkle 7466 LAM!, AHFH!; Thome 37526
RSA!; Philbrick & McPherson B68-221 RSA!; SBBG.

Santa Catalina: Fosberg S4533 LAM!; Thome 35748 RSA!;
Thome 42845 RSA!; SBBG.

San Clemente: Dunkle 7306 LAM!, AHFH!; Munz 6665
POM!; Raven 17135 RSA!; SBBG.

Guadalupe: Palmer 20 CM!; Wiggins & Ernst 21 DS!; Nor-
ris s.n. May 1, 1951 LA!.

Erodium obtusiplicatum (Maire, Weiller & Wilcz.) J.T. How-
ell

Santa Rosa: Munz & Crow 1 1636 LA!, POM!; Thome et
al. 48874b RSA!; Thome et al. 49008 RSA!.

Santa Catalina: Thome, Rollins, Propst & Carolin 36750
RSA!; Thome, Rollins, Propst & Carolin 36757 RSA!.

Erodium texanum Gray

Santa Catalina: Trask s.n. in Mar. 1901 LAM!.

Geranium carolinianum L.

Santa Rosa: SBBG; SBM.

Santa Cmz: SBM.

Santa Catalina: Fosberg S4319 LAM!; Fosberg S4593
LAM!; Fosberg S4627 LAM!; RSA-POM.

Geranium dissectum L.

Santa Rosa: Thome et al. 48750 RSA!.

Pelargonium x hortorum Bailey

Santa Rosa: SBBG.

Santa Cmz: SBBG.

San Nicolas: Wier & Beauchamp s.n. Jun. 30, 1978
RSA#289077!.

Santa Catalina: Thome 36477 RSA!; SBBG.

Pelargonium peltatum (L.) L’Her. ex Ait.

San Nicolas: Foreman 106 UC!; Wier & Beauchamp s.n.
Jun. 30, 1978 RSA!.

Hydrophyllaceae

Emmenanthe penduliflora Benth.

Santa Cmz: Hoffmann s.n. Apr. 22, 1932 POM!; SBM.

Santa Catalina: Fosberg S4694 LAM!; Dunkle 2446
AHFH!; Thome 35972 RSA!; SBBG.

San Clemente: Dunkle 7302 LAM!, AHFH!, RSA!; Raven
17722 RSA!; SBBG.

Guadalupe: Palmer 73 CM!; Carlquist 448 RSA!; Moran
5962 RSA!.

Eriodictyon traskiae Eastw. ssp. traskiae

Santa Catalina: Trask s.n. in Mar. 1902 LAM!; Fosberg
S4790 LAM!; Dunkle 1 96 1 AHFH!, POM!; CAS; SBBG.

68 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Eucrypta chrysanthemifolia (Benth.) Greene var. chrysan-
themifolia

San Miguel: Munz & Norris 11783 POM!; SBBG; SBM.
Santa Rosa: Thome et al. 48807 RSA!; Blakley & Smith
3082 RSA!; SBBG; SBM.

Santa Cruz: Fosberg 7631 LAM!, LA!; POM!; Munz &
Crow 11524 POM!; Wolf 28 17 RSA!; SBBG; SBM.
Anacapa: Moran 725 LAM!; SBBG; SBM.

Santa Catalina: Fosberg S4723 LAM!; Fosberg S4412
LAM!; Moran 683 LAM!; RSA-POM; SBBG.

San Clemente: Thorne 36111 RSA!; Raven 17695 RSA!;
Raven 17736 RSA!; SBBG.

Guadalupe: Anthony 252 POM!; Moran 5684 RSA!; Flow-
ell 8322 POM!.

Nemophila menziesii H. & A. ssp. menziesii
Santa Catalina: Moran 681 LAM!, DS!, RSA!; SBBG.
Nemophila pedunculata Dougl. ex Benth.

San Miguel: Hoffmann s.n. Apr. 10, 1930 SBM#9226!,
CAS#1 78505!.

Santa Rosa: Hoffmann s.n. Apr. 17, 1929 SBM#5992!;
Hoffmann s.n. Apr. 17, 1929 SBM#5994!; Raven 14934
RSA!; SBBG.

Santa Cruz: Munz & Crow 1 1539 POM!; Hoffmann s.n.

in Mar. 1929 CAS# 1683 19!; SBBG.

San Nicolas: (Howell 1935: Howell 8213 at CAS).
Phacelia cicutaria Greene ssp. hispida (Gray) Beauchamp ex
Thome

Santa Rosa: SBM.

Santa Cruz: Fosberg 7538 POM!; Thome & Everett 36830
RSA!; Raven & Smith 15263 RSA!; SBBG; SBM.
Anacapa: SBBG; SBM.

Santa Catalina: Fosberg S4432 LAM!; Fosberg S4336
LAM!; Trask s.n. in Mar. 1901 ND-G#042100!; RSA-
POM; SBBG.

Phacelia cinerea Eastw. ex Macbr.

San Nicolas: Trask s.n. in Apr. 1901 LAM!, ND-
G#042098!.

Phacelia distans Benth.

San Miguel: Dunkle 8407 LAM!, AHFH!; Munz & Crow
11813 POM!; Greene s.n. in Sep. 1 886 CAS#938!; SBBG;
SBM.

Santa Rosa: Dunkle 8476 AHFH!; Moran 795 LAM!, RSA!;

Munz & Crow 1 1753 LA!, POM!; SBBG; SBM.

Santa Cruz: Fosberg 7538 LAM!; Wolf 2794 RSA!; Web-
ster et al. 96 RSA!; SBBG; SBM.

Anacapa: Moran 721 LAM!, RSA!; Howell 3801 RSA!;
SBBG; SBM.

Santa Barbara: Dunkle 7460 LAM!; Dunkle 7441 LAM!,
AHFH!; Dunkle 8122 LAM!, AHFH!; SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Fosberg
S4598 LAM!; RSA-POM; SBBG.

San Clemente: Thome 35989 RSA!; Munz 6738 POM!;
Raven 17306 RSA!; SBBG.

Phacelia divaricata (Benth.) Gray var. insu/aris (Munz) Munz
San Miguel: Munz & Norris 1 1829 POM!; Hoffmann s.n.
Apr. 20, 1932 POM!; SBBG; SBM.

Santa Rosa: Moran 800 LAM!, RSA!; Munz 11756 POM!;
SBBG; SBM.

Phacelia floribunda Greene

San Clemente: Dunkle 72 1 2 LAM!, AHFH!, RSA!; Thorne
36112 RSA!; Raven 17243 RSA!; SBBG; SBM.
Guadalupe: Anthony 242 LAM!; Moran 6712 LA!, RSA!;
Moran 5653 RSA!.

Phacelia grandiflora (Benth.) Gray
Santa Rosa: SBM.

Santa Cruz: SBBG; SBM.

Santa Catalina: Raven 17859 RSA!; SBBG.

Phacelia lyonii Gray

Santa Catalina: Davidson, A. s.n. in Jun. 1897 LAM!;
Dunkle 2 1 46 LAM!, AHFH!; Fosberg S4749 LAM!; LA;
RSA-POM; SBBG.

San Clemente: Moran 682 RSA!; Raven 17690 RSA!;
Thorne 36045 RSA!; SBBG.

Phacelia phyllomanica Gray

Guadalupe: Franceschi 43 LAM!, RSA!; Moran 7836 RSA!;
Moran 13782 RSA!.

Phacelia ramosissima Dougl. ex Lehm. var. austrolit oralis
Munz

Santa Rosa: Dunkle 8525 AHFH!; Thorne et al. 48779
RSA!; Thorne et al. 48751 RSA!.

Santa Cruz: Greene s.n. in Jul.-Aug. 1 886 ND-G#042054!;
Greene s.n. in Jul.-Aug. 1886 ND-G#042053!; Mower
s.n. Aug. 22, 1966 LA#96261!; RSA-POM.

Phacelia ramosissima Dougl. ex Lehm. var. montereyensis
Munz

San Miguel: SBM.

Santa Rosa: Dunkle 8476 LAM!; Dunn, N. s.n. May 15,
1932 LA!; SBM.

Phacelia viscida (Benth.) Torn.

Santa Rosa: Fosberg 7516 LAM!; Fosberg 7638 LAM!;
Blakley 3150 RSA!; SBBG.

Santa Cruz: Fosberg 7521 LAM!; Moran 757 LAM!, RSA!;

Raven & Smith 15206 RSA!; LA; SBBG; SBM.
Anacapa: SBBG.

Santa Catalina: Trask s.n. in Mar. 1900 NY!; Trask s.n.
in Mar. 1901 ND-G#04072!, NY!, US!.

Pholistoma auritum (Lindl.) Lilja ex Lindl.

Santa Cruz: Fosberg 7664 LAM!, LA!, POM!; Elmore s.n.
Apr. 18, 1936 AHFH!; Raven & Smith 15153 RSA!;
SBBG; SBM.

Santa Barbara: SBBG.

Santa Catalina: Dunkle 2089 LAM!, AHFH!; Moran 684
LAM!, RSA!; Fosberg S4592 LAM!, POM!; SBBG; SBM.
San Clemente: Dunkle 7238 LAM!, AHFH!, RSA!; Thome
42869 RSA!; Raven 17346 RSA!; SBBG.

Pholistoma racemosum (Nutt.) Const.

Santa Cruz: Fosberg 7667 LAM!, LA!; Elmore s.n. Apr.

18, 1936 USC!; Wolf 2900 RSA!; SBBG; SBM.

Santa Barbara: Dunkle 7443 LAM!, AHFH!; Thome 375 1 3
RSA!; Blakley 5651 RSA!; SBBG.

Santa Catalina: Trask s.n. in Mar. 1 900 LAM!; Grant 367 1
LAM!; Millspaugh 4687 F!; LA; RSA-POM; SBBG; SBM.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 69

San Clemente: Dunkle 7219 LAM!, AHFH!; Moran 574
LAM!; Elmore 412 AHFH!; RSA-POM; SBBG.
Guadalupe: Anthony 254 RSA!; Moran 5686 RSA!.
Juglandaceae
Juglans californica Wats.

Santa Catalina: Johnstone s.n. May 20, 1934 USC!; Fos-
berg S4848 LAM!; Thome & Everett 34674 RSA!; SBBG.
Lamiaceae

Lamium amplexicaule L.

Santa Cruz: Wolf 2886 RSA!; SBBG; SBM.

Lepechinia calycina (Benth.) Epl. in Munz
Santa Rosa: Moran 790 LAM!, RSA!; SBBG.

Santa Barbara: SBBG.

Lepechinia fragrans (Greene) Epl.

Santa Rosa: Epling & Erickson s.n. Aug. 8, 1937 LA!;

Thome et al. 48997 RSA!; Raven 15008 RSA!; SBM.
Santa Cruz: Greene s.n. in Jul.-Aug. 1886 CAS#1140!;
Clokey 5044 POM!; Jones s.n. Mar. 25, 1929 POM!,
RSA!; LA; SBM.

Santa Catalina: Trask s.n. in May 1901 LAM!; Fosberg
S4786 LAM!, POM!; Thome 36329 LA!, RSA!.
Marrubium vulgare L.

San Miguel: Dunkle 8374 LAM!, AHFH!; SBBG; SBM.
Santa Rosa: SBBG; SBM.

Santa Cruz: Dunkle 8638 LAM!, AHFH!; Clokey 5043
LA!, POM!; SBBG; SBM.

San Nicolas: Newman 1 14 Pacific Missile Test Center, Pt.
Mugu!

Santa Catalina: Thome & Everett 34428 RSA!; Fosberg
S4459 LAM!; Dunkle 1874 AHFH!; SBBG.

San Clemente: DeBuhr & Wallace 71 1 LAM!; House &
Grumbles s.n. Aug. 5-13, 1930 USC!; Elmore 417
AHFH!; RSA-POM; SBBG.

Mentha citrata Ehrh.

Santa Catalina: Thome 36521 RSA!; Thome & Everett
34957 RSA!; SBBG.

Mentha spicata L.

Santa Catalina: Thome 36637 RSA!.

Nepeta cataria L.

Santa Catalina: Millspaugh 4894 F!; Nuttall 831 F!; Pen-
dleton 1391 POM!.

Pogogyne tenuiflora Gray
Guadalupe: Palmer 65 NY!.

Salvia apiana Jeps. var. apiana
Santa Catalina: Fosberg S4800 LAM!; Fosberg 4808 LAM!,
POM!; Eaton s.n. Jul. 15, 1956 LAM!; SBBG; SBM.
Salvia brandegei Munz

Santa Rosa: Dunkle 8427 LAM!, AHFH!; Epling s.n. Apr.
20, 1940 LA!; Thome et al. 48929 RSA!; SBBG; SBM.
Salvia columbariae Benth.

Santa Rosa: SBM.

Santa Cruz: SBBG; SBM.

Santa Catalina: Dunkle 1774 AHFH!, POM!; Fosberg
S4699 LAM!, POM!; Thome 35945 RSA!.

San Clemente: Munz 6478 POM!; Thome 42830 RSA!;
Raven 17688 RSA!.

Salvia mellifera Greene
Santa Rosa: SBBG; SBM.

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 LAM!; Fosberg
7707 LAM!, POM!; Raven & Smith 15255 RSA!; SBBG;
SBM.

Anacapa: Dunkle 7635 LAM!; Moran 736 LAM!, RSA!;
SBBG; SBM.

Santa Catalina: Fosberg S4291 LAM!; Fosberg S4805
LAM!; Dunkle 1725 AHFH!, POM!; RSA; SBBG; SBM.
Satureja douglasii (Benth.) Briq.

Santa Catalina: Trask s.n. in Aug. 1902 LAM!; Fosberg
S4702 LAM!, POM!; Dunkle 2008 AHFH!, POM!;
SBBG.

Satureja palmeri (Gray) Briq.

Guadalupe: Palmer 66 NY!, CM!.

Scutellaria tuberosa Benth. ssp. australis Epl.

Santa Cruz: Fosberg 7694 LAM!, LA!; Fosberg 7659 LAM!,
LA!; Moran 749 LAM!; RSA-POM; SBBG; SBM.
Stachys bullata Benth.

Santa Rosa: Dunkle 8530 LAM!; Dunn, N. s.n. May 15,
1932 LA!; Munz & Crow 1 1589 LA!; SBBG; SBM.
Santa Cruz: Hoffmann 218 LA!; Abrams & Wiggins 59
NY!; Eastwood 6414 US!; RSA-POM; SBBG; SBM.
Anacapa: Dunkle 7659 LAM!; Moran 737 LAM!; SBBG;
SBM.

Trichostema lanceolatum Benth.

Santa Catalina: Dunkle 2 1 50 LAM!, AHFH!; Thome 36707
RSA!; Thome, Propst & Hoefs 45111 RSA!; SBBG.
Linaceae

Hesperolinon micranthum (Gray) Small
Santa Catalina: RSA-POM; SBBG.

Loasaceae

Mentzelia affinis Greene
Santa Cruz: Fosberg 7536 LAM!, POM!; SBM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!.

San Clemente: SBBG; SBM.

Mentzelia micrantha (H. & A.) T. & G.

Santa Cruz: Clokey 5028 NY!; Clokey 5008 POM!; SBBG;
SBM.

Santa Catalina: Dunkle 2465 LAM!; Heller 8954 NY!;
Trask s.n. in Mar. 1896 MO!; RSA-POM; SBBG; SBM;
US.

San Clemente: Munz 6749 POM!; SBBG.

Guadalupe: Palmer 893 ND-G!, US!; Anthony 259 US!;
Wiggins & Ernst 63 US!.

Lythraceae

Ammannia coccinea Rottb.

Santa Catalina: Thome 36710 RSA!.

Lythrum californicum T. & G.

Santa Cruz: Greene s.n. in Jul.-Aug. 1886 ND-G#034589!.
Malvaceae

Althaea rosea (L.) Cav.

Santa Catalina: Thome 36475 RSA!.

Eremalche exilis (Gray) Greene
Santa Catalina: Trask s.n. in Mar. 1901 LAM!, NY!.

San Clemente: Trask 218 NY!.

70 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Lavatera assurgentiflora Kell.

San Miguel: Greene s.n. in Sep. 1886 MO!, POM!; SBBG.
Santa Rosa: Dunkle 8529 LAM!; Elmore 191 AHFH!;
SBBG.

Santa Cruz: RSA-POM.

Anacapa: (Philbrick 1980: Timbrook & Philbrick 652 and
653 at SBBG).

San Nicolas: Wier & Beauchamp s.n. Jul. 4, 1978 RSA!,
UC!.

Santa Catalina: Fosberg 7146 LAM!, POM!; Trask s.n. in
Mar. 1896 MO!; Moran 625 LAM!, RSA!; SBBG; SBM.
San Clemente: Raven 17303 LA!, RSA!; Blakley 6414 RSA!;
Raven 17579 RSA!.

Lavatera linsayi Moran

Guadalupe: Lindsay 1812 DS!; Moran 15118a LAM!;
Moran 2630 RSA!.

Lavatera occidentals Wats.

Guadalupe: Franceschi 12 LAM!; Anthony 247 MO!;
Palmer 17 MO!, CM!.

Malacothamnus clementinus (M. & J.) Kearn.

San Clemente: Munz 6684 POM! CAS!, DS!; Thome 42778
RSA!; Raven 17977 RSA!; SBBG.

Malacothamnus fasciculatus (Nutt.) Greene ssp. catalinensis
(Eastw.) Thome

Santa Catalina: Fosberg S5427 LAM!, POM!; Wallace &
Haefs 1407 RSA!; Dunkle 1932 LAM!; RSA-POM;
SBBG; SBM.

Malacothamnus fasciculatus (Nutt.) Greene var. nesioticus
(Rob.) Keam.

Santa Cruz: Hoffmann s.n. Jun. 28, 1930 POM!; Greene
s.n. in Jul.-Aug. 1886 CAS#743!; SBBG; SBM.

Malva parviflora L.

San Miguel: Dunkle 8393 LAM!, AHFH!; SBBG; SBM.
Santa Rosa: Thome et al. 488 1 9 RSA!; Thorne et al. 48879
RSA!; Raven, Blakley & Omduff 14875 RSA!; SBBG;
SBM.

Santa Cruz: Dunkle 8632 LAM!, AHFH!; Hoffman s.n.
Apr. 12,1931 LAM!; Hoffmann s.n. Sep. 2 1 , 1 930 POM!;
SBBG; SBM.

Anacapa: SBBG.

San Nicolas: Kanakoff s.n. Apr. 26, 1940 LAM!; Dunkle
8326 LAM!, AHFH!; Foreman, Evans & Rainey 1 1 RSA!;
SBBG; SBM.

Santa Barbara: Dunkle 8117 LAM!; Bryan, Dr. & Mrs. s.n.

Jul. 14, 1922 LAM!; Thome 37509 RSA!; SBBG; SBM.
Santa Catalina: Fosberg S4472 LAM!; Fosberg S5434
LAM!; Dunkle 1856 AHFH!; RSA-POM; SBBG.

San Clemente: Dunkle 7293 LAM!; Munz 6784 POM!;

Raven 17129 RSA!; SBBG.

Guadalupe: Howell 826! POM!.

Malvella leprosa (Ortega) Krapovickas

Santa Catalina: Blakley 5378 LAM!, CAS!, RSA!; SBBG.
San Clemente: Trask 258 US!; Munz 6622a POM!; Raven
17956 RSA!, CAS!.

Guadalupe: (Clement 1957: Thobum, Greene & Wing s.n.
in Jun. 1897 at DS).

Sidalcea malvaeflora (DC.) Gray ex Benth. ssp. malvaeflora
San Miguel: Munz & Norris 1 1782 POM!; SBBG; SBM.
Santa Rosa: Moran 780 LAM!, MO!; Elmore 181 AHFH!;
Dunn, N. s.n. May 15, 1932 LA!; RSA-POM; SBBG;
SBM.

Santa Cruz: Raven & Smith 15300 RSA!; Webster et al.
95 RSA!; SBBG.

Sphaeralcea palmeri Rose

Guadalupe: Rempel 759-37 LAM!; Carlquist 469 RSA!;
Moran 17332 RSA!.

Sphaeralcea sulphur ea Wats.

Guadalupe: Palmer 18 CM!; Franceschi 13 RSA!.
Moraceae
Ficus carica L.

San Miguel: SBBG.

Santa Cruz: SBBG; SCIR.

Santa Catalina: RSA-POM; SBBG.

Myrtaceae

Eucalyptus globulus Labill.

Santa Rosa: SBBG.

Santa Cruz: SBBG.

Anacapa: SBBG.

San Nicolas: Wier & Beauchamp s.n. Jul. 4, 1978 RSA!;
SBBG.

Santa Catalina: RSA-POM.

Nyctaginaceae
Abronia latifolia Esch.

San Miguel: Munz & Fosberg 1 1792 POM!; SBM.
Abronia maritima Nutt, ex Wats.

San Miguel: Dunkle 8390 LAM!, AHFH!, RSA!; Elmore
330 LAM!, AHFH!; Elmore 340 AHFH!; SBBG; SBM.
Santa Rosa: Munz & Crow 1 1567 POM!; Raven 14997
RSA!; Thome et al. 49022 RSA!; SBM.

Santa Cruz: Dunkle 8583 LAM!, RSA!; Balls & Blakley
23637 RSA!; Balls & Blakley 23694 RSA!; SBM.
Anacapa: Elmore 243 AHFH!; SBM.

San Nicolas: Dunkle 8342 LAM!, RSA!; Foreman & Lloyd
139 RSA!; Raven & Thompson 20703 RSA!; SBBG.
Santa Catalina: Fosberg S488 1 LAM!, POM!; Dunkle 1 892
AHFH!, POM!; Moran 675 LAM!, RSA!; SBBG; SBM.
San Clemente: Dunkle 7278 LAM!, AHFH!; Elmore 381
AHFH!; Thome 42933 RSA!; SBBG.

Abronia umbel/ata Lam.

San Miguel: Dunkle 8373 LAM!, AHFH!; Elmore 330a
AHFH!; Elmore 339 AHFH!; RSA-POM; SBBG; SBM.
Santa Rosa: Dunkle 8490 LAM!; Moran 792 LAM!; Epling
& Erickson s.n. Aug. 8, 1937 LA!; RSA-POM; SBBG;
SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

San Nicolas: Trask 23 LAM!; Kanakoff s.n. Apr. 14, 1940
LAM!; Dunkle 8316 LAM!; LA; RSA-POM; SBBG.
Santa Catalina: Thome 39383 RSA!; Thome & Everett
34934 RSA!; SBBG.

San Clemente: Dunkle 7266 LAM!; Elmore s.n. Nov. 23,
1939 AHFH!; Raven 17297 RSA!.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 71

Mirabilis californica Gray var. californica

Santa Cruz: Clokey 4923 LAM!; Fosberg 7518 LAM!;

Howell 6287 US!; RSA-POM; SBBG; SBM.

Anacapa: Dunkle 7671 LAM!, AHFH!, RSA!; SBBG; SBM.
Santa Barbara: Dunkle 7433 LAM!; Dunkle 8107 LAM!;
Thome 37483 RSA!.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!, NY!, US!;
Moran 669 LAM!; Eastwood 6443 LA!; RSA-POM;
SBM.

San Clemente: Dunkle 7234 LAM!; Dunkle 7235 AHFH!,
RSA!; Thome 42871 RSA!; SBBG.

Mirabilis heimerlii (Standi.) Macbr.

Guadalupe: Palmer 886 ND-G!, NY!; Rose 16021 NY!;
Carlquist 481 RSA!; LAM.

Oleaceae

Hesperelaea palmeri Gray

Guadalupe: Palmer 81 NY!, CM!.

Olea enropaea L.

Santa Cruz: SBBG.

Onagraceae

Camissonia californica (Nutt. ex. T. & G.) Raven
Santa Rosa: SBM.

Santa Cruz: Breedlove 28 1 6 RSA!; Thome & Everett 36842
RSA!; Raven & Smith 15274 RSA!; SBBG; SBM.

Santa Catalina: Dunkle 1943 AHFH!, POM!.

Camissonia cheiranthifolia (Homem. ex Spreng.) Raim. in
Engl. & Prantl ssp. cheiranthifolia
San Miguel: Dunkle 8379 LAM!, AHFH!; Dunkle 8414
AHFH!; Elmore 313 AHFH!; RSA-POM; SBBG; SBM.
Santa Rosa: Moran 791 LAM!, RSA!; Dunkle 8467 LAM!,
AHFH!; Thome et al. 49021 RSA!; SBBG; SBM.

Santa Cruz: Raven & Smith 15312 RSA!; Webster et al.
89 RSA!; SBBG; SBM.

San Nicolas: Dunkle 8309 LAM!, AHFH!; Kanakoffs.n.
Apr. 18, 1940 LAM!; Foreman, Evans & Rainey 79 LA!,
RSA!; SBBG.

Santa Barbara: Hemphill s.n. UC# 172325!.

San Clemente: Blakley 5255 RSA!; Raven 17958 RSA!;
Raven 17627 RSA!; SBBG.

Camissonia cheiranthifolia (Homem. ex Spreng.) Raim. in
Engl. & Prantl ssp. sujfruticosa (Wats.) Raven
San Nicolas: Trask s.n. in Apr. 1901 LAM!; Raven &
Thompson 20761 LA!, RSA!; SBM.

Camissonia guadalupensis (Wats.) Raven ssp. Clementina
(Raven) Raven

San Clemente: Thome 35991 RSA!; Raven 17125 RSA!,
DS!; Raven 186760 RSA!; SBBG.

Camissonia guadalupensis (Wats.) Raven ssp. guadalupensis
Guadalupe: Moran & Ernst 6737 DS!.

Camissonia hirtella (Greene) Raven
Santa Cruz: Clokey 5010 POM!.

Camissonia ignota (Jeps.) Raven
Santa Cruz: Hoffmann s.n. Mar. 24, 1929 POM!.
Camissonia intermedia Raven
Santa Cruz: Moran 764 LAM!; Hoffmann s.n. Mar. 25,
1932 POM!; Raven & Smith 15197 RSA!.

Santa Catalina: Trask s.n. in May 1896 F!; Knopf 428 F!;
Raven 18171 RSA!.

Camissonia micrantha (Homem. ex Spreng.) Raven
San Miguel: SBBG; SBM.

Santa Rosa: Munz & Crow 1 1694 LA!, POM!; Hoffmann
s.n. Apr. 16, 1929 POM!; Thome et al. 49037 RSA!;
SBBG; SBM.

Santa Cruz: Jones s.n. Mar. 25, 1929 POM!; Thome &
Everett 36819 RSA!; SBBG; SBM.

Santa Catalina: Fosberg S4315 LAM!; Thome 36501 RSA!;
SBM.

San Clemente: Piehl 62387 RSA!; SBBG.

Camissonia robusta Raven
San Miguel: SBBG.

Santa Cruz: SBBG.

Santa Catalina: Trask s.n. in May 1897 LAM!; Dunkle
1937 AHFH!; Raven 17811 RSA!.

San Clemente: Dunkle 7298 LAM!, AHFH!, RSA!; Munz
6743 POM!; SBBG.

Guadalupe: Copp 151 RSA!; Moran 2891 RSA!; Moran
5661 RSA!.

Camissonia strigulosa (F. & M.) Raven
Santa Rosa: Moran 801 LAM!; Munz & Hoffman 11742
POM!; SBBG; SBM.

Clarkia davyi (Jeps.) Lewis & Lewis
Santa Rosa: Raven 15003 LA!, RSA!; Raven 15007 RSA!;
Raven 14955 RSA!; LA; SBBG.

Clarkia epilobioides (Nutt.) Nels. & Macbr.

Santa Rosa: Blakley & Smith 3089 RSA!; Thome et al.
48773 RSA!; SBBG; SBM.

Santa Cruz: Elmore s.n. Apr. 18, 1936 AHFH!, RSA!,
USC!; Munz & Crow 11861 POM!; Raven & Smith
15151 RSA!; SBBG.

Santa Catalina: Haefs 201 MO!; Fosberg 8117 LAM!,
POM!; Smith 5965 US!; LA; NY; RSA-POM; SBBG.
San Clemente: Trask 172 US!; Trask s.n. in May 1895 US!;
Raven 17710 RSA!; SBBG.

Clarkia prostrata Lewis & Lewis
Santa Rosa: Youngberg s.n. May 29, 1938 POM!; Hoff-
mann 692 POM!; Hoffmann s.n. Jul. 7, 1930 POM!.
Clarkia purpurea (Curt.) Nels. & Macbr. ssp. quadrivulnera
(Dougl. in Lindl.) Lewis & Lewis
Santa Rosa: SBM.

Santa Cruz: Sauer et al. 5498 RSA!.

Santa Catalina: Fosberg S4696 LAM!; POM!; Fosberg
S4868 LAM!; Haefs & Propst 220 MO!; RSA-POM;
SBBG.

Clarkia unguiculata Lindl.

Santa Cruz: SBM.

Santa Catalina: Dunkle 2464 AHFH!.

Epilobium canum (Greene) Raven ssp. canum

Santa Rosa: Epling & Erickson s.n. Aug. 8, 1937 LA!; RSA-
POM; SBBG; SBM.

Santa Cruz: Greene s.n. in Jul. -Aug. 1886 MO!; Hoffmann
s.n. Sep. 20, 1930 LA!; RSA-POM; SBBG; SBM.
Anacapa: RSA-POM; SBBG; SBM.

Santa Catalina: Fosberg S5419 MO!; Ewan 10804 LA!,
MO!; Nuttall 660 MO!; RSA-POM; SBBG.

San Clemente: Trask s.n. in Dec. 1896 MO!; Moran et al.
22691 MO!; RSA-POM; SBBG.

72 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Epilobium ciliatum Raf. ssp. ciliatum
San Miguel: Dunkle 8419 LAM!, RSA!.

Santa Cruz: Hoffmann s.n. Sep. 21, 1931 LAM!; Hoffmann
s.n. Jul. 1, 1930 POM!; Balls & Blakley 23722 RSA!;
SBBG; SBM.

Santa Catalina: Fosberg S5370 LAM!; Trask s.n. in Aug.
1900 LAM!; Thorne 36634 RSA!; SBBG; SBM.
Epilobium foliosum (T. & G.) Suksd.

Guadalupe: Palmer 31 MO!.

Gaura sinuata Nutt, ex Ser. in DC.

Santa Catalina: Thome 36478 RSA!; SBBG.

Ludwigia peploides (HBK.) Raven ssp. peploides

Santa Cruz: Dunkle 8569 LAM!, AHFH!; Balls & Blakley
23680 RSA!.

Oenothera elata HBK. ssp. hirsutissima (Gray ex Wats.) Die-
trich

Santa Cruz: Stanton s.n. in Nov. 1967 RSA!; Daily 171
SCIR!.

Orobanchaceae
Orobanche bulbosa G. Beck

Santa Rosa: Hoffmann 155 POM!; Thome et al. 48963
RSA!; Hoffmann s.n. May 9, 1932 POM!; SBM.

Santa Cruz: Hoffmann s.n. Jun. 15, 1930 POM!; SBBG;
SBM.

Santa Catalina: Fosberg S4687 LAM!, POM!; Trask s.n.
in Apr. 1900 US!; Dunkle 2449 AHFH!; SBBG.
Orobanche californica Cham. & Schlecht. ssp. grandis Heck-
ard

Santa Rosa: Hoffmann s.n. Jun. 11, 1930 SBM.
Orobanche fasciculata Nutt.

Santa Rosa: Hoffmann s.n. May 8, 1932 POM!; SBM.
Santa Cruz: Daily 376 SCIR!.

Santa Catalina: Pendleton 1353 POM!; Thome 36235 RSA!;
SBBG.

Orobanche parishii (Jeps.) Heckard ssp. brachyloba Heckard
San Miguel: SBBG.

Santa Rosa: Hoffmann s.n. Aug. 9, 1931 SBM!.

Santa Cmz: Daily 530 SCIR!; SBBG.

San Nicolas: Raven & Thompson 20794 RSA!; Wier &
Beauchamp s.n. Jun. 31, 1978 RSA!; Wier & Beauchamp
s.n. Jul. 1, 1978 RSA!; SBBG.

Santa Catalina: RSA-POM; SBBG.

Orobanche uniflora L. ssp. occidentalis (Greene) Abrams ex
Ferris

Santa Cruz: Pierson 11088 RSA!; Harvey s.n. Apr. 20,
1936 POM#223655!; SBM.

Oxalidaceae

Oxalis albicans HBK. ssp. californica (Abrams) Eiten
Santa Cmz: Dunkle 8544 LAM!, AHFH!; Fosberg 7672
LAM!; Thome & Everett 36841 RSA!; SBBG; SBM.
Santa Catalina: Fosberg S4586 LAM!, POM!; Trask s.n.
in Apr. 1902 LAM!; Dunkle 2866 AHFH!; LA.

Oxalis albicans HBK. ssp. pilosa (Nutt.) Eiten
Santa Cruz: SBM.

Oxalis corniculata L.

Santa Cmz: SBM.

Santa Catalina: Thome 36428 RSA!; Thome 36331 RSA!;
SBBG.

Oxalis pes-caprae L.

Santa Catalina: Fosberg S4523 LAM!; Thorne & Everett
34986 RSA!; SBBG.

San Clemente: Elmore 423 AHFH!.

Papaveraceae

Dendromecon rigida Benth. ssp. harfordii (Kell.) Raven
Santa Rosa: Yates s.n. LAM#19889!; Dunkle 8497 LAM!,
AHFH!; Harford s.n. in 1873 CAS#2405!; LA; RSA-
POM; SBBG.

Santa Cmz: Clokey 4942 LAM!; Fosberg 7605 LAM!, LA!,
POM!; Greene s.n. in Jul. -Aug. 1886 CAS#2403!,
DS#97726!; SBBG.

Dendromecon rigida Benth. ssp. rhamnoides (Greene) Thome
Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Fosberg
S4580 LAM!; Moran 613 LAM!; LA; RSA-POM.

San Clemente: Trask s.n. in Jun. 1903 US!.

Eschscholzia californica Cham. var. californica

San Miguel: Dunkle 8385 LAM!, AHFH!, NY!; Moran
3447 POM!; Munz & Norris 11772 POM!.

Santa Rosa: Dunkle 8456 LAM!; Elmore 187 AHFH!;

Thome et al. 48873 RSA!; LA; SBM.

Santa Cruz; Fosberg 7640 LAM!; Dunkle 8361 LAM!,
AHFH!, RSA!; Greene s.n. in Jul.-Aug. 1 886 CAS#299!,
US!; LA; SBBG; SBM.

Santa Catalina: Brandegee s.n. May 17-25, 1916 LAM!;

Moran 612 LAM!; Fosberg S4954 LAM!; SBBG.
Guadalupe: Greene s.n. Apr. 23, 1885 NY!.

Eschscholzia californica Cham. var. maritima (Greene) Jeps.
San Miguel: Yates s.n. LAM# 1 9879!; Elmore 320 AHFH!;

Elmore 327 AHFH!; RSA-POM; SBBG.

Santa Rosa: SBBG; SBM.

Santa Cmz: Hoffmann s.n. Sep. 21, 1930 POM!; SBBG;
SBM.

Eschscholzia californica Cham. var. peninsularis (Greene)
Munz

Santa Cruz: Pierson 1 1066 RSA!; Wolf 2863 RSA!; Wolf
2792 RSA!.

Santa Catalina: Fosberg S4650 LAM!; Millspaugh 4854 F!;
Thome & Everett 34501 RSA!.

Eschscholzia elegans Greene

Guadalupe: Palmer 3 (in part) NY!, (in part) US!, CM!;
Greene s.n. Apr. 23, 1885 CAS#288!, NY!; Ernst 273
US!.

Eschscholzia frutescens (Greene) J.T. Howell

Guadalupe: Franceschi s.n. in Jan. 1893 CAS#2654!.
Eschscholzia palmeri Rose

Guadalupe: Rempel 759-37 LAM!; Palmer 889 NY!, US!;
Howell 8176 NY!, POM!, US!; Ernst 272 US!.
Eschscholzia ramosa (Greene) Greene
Santa Rosa: SBM.

Santa Cmz: SBBG; SBM.

Santa Barbara: RSA-POM; SBBG.

Santa Catalina: Brandegee, K. s.n. May 17-25, 1916 LAM!;
Fosberg S4598 LAM!; Trask 307 NY!, US!: RSA-POM;
SBBG.

San Clemente: Dunkle 7213 LAM!, AHFH!; Dunkle 7301
LAM!, AHFH!; Trask 263 NY!, US!; RSA-POM; SBBG.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 73

Guadalupe: Anthony 231 LAM!, POM!; Rose 16032 NY!;
Franceschi 20 SBM!; CAS; LA; RSA-POM; US.
Meconella denticulata Greene

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 LAM!; RSA-
POM; SBBG; SBM.

Papaver californicum Gray

Santa Cruz: RSA-POM; SBBG; SBM.

Papaver somniferum L.

Santa Catalina: RSA-POM.

Platystemon californicus Benth.

San Miguel: RSA-POM; SBBG; SBM.

Santa Rosa: Brandegee s.n. in Jun. 1888 CAS#977!; RSA-
POM; SBBG; SBM.

Santa Cruz: Moran 765 LAM!; Elmore 458 AHFH!; Fos-
berg 7555 LAM!, LA!; RSA-POM; SBBG; SBM.
Anacapa: SBBG; SBM.

San Nicolas: Dunkle 8332 LAM!; Kanakoff s.n. LAM!;

Trask 26 LAM!; LA; RSA-POM; SBBG; SBM.

Santa Barbara: Dunkle 7400 LAM!; Trask 1 1 CAS!; Trask
s.n. in May 1901 CAS#969!; RSA-POM; SBBG.

Santa Catalina: Brandegee, T.S. s.n. in May 1890 UC!;

Trask s.n. in Mar. 1897 CAS#966!.

Guadalupe: Brandegee, T.S. s.n. Mar. 20, 1897 UC!.
Romneya coulteri Harv.

Santa Catalina: RSA-POM; SBBG.

Stylomecon heterophylla (Benth.) G. Taylor
San Miguel: SBBG; SBM.

Santa Rosa: Munz & Crow 1 1652 LA!; RSA-POM; SBBG;
SBM.

Santa Cruz: Fosberg 7620 LA!; RSA-POM; SBBG; SBM.
Anacapa: SBBG.

Santa Barbara: RSA-POM; SBBG; SBM.

Santa Catalina: RSA-POM; SBBG.

San Clemente: RSA-POM; SBBG.

Pittosporaceae

Solly a heterophylla Lindl.

Santa Catalina: Nuttall 801 F!.

Plantaginaceae

Plantago bigelovii Gray ssp. californica (Greene) Bassett
San Miguel: RSA-POM; SBM.

Santa Rosa: RSA-POM; SBM.

Plantago coronopus L.

Santa Catalina: Trask 308 US!; Johnson 1758 US!; Fosberg
4671 LAM!, US!, MO!; Davidson, A. s.n. Jun. 26, 1891
LAM!; RSA-POM; SBBG.

Plantago erecta Morris ssp. erecta
Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7683 LAM!, LA!; RSA-POM; SBBG;
SBM.

Anacapa: SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!, NY!, US!;
Fosberg S4548 LAM!, NY!; Millspaugh 4910 F!; LA;
RSA-POM; SBBG.

San Clemente: Dunkle 7324 LAM!, AHFH!; RSA-POM;
SBBG.

Plantago hirtella HBK. ssp. galeottiana (Dene.) Thome
Santa Rosa: RSA-POM; SBM.

Santa Cruz: RSA-POM; SBM.

Plantago lanceolata L.

Santa Cruz: Daily 316 SCIR!.

San Clemente: RSA-POM.

Plantago major L.

Santa Cruz: Dunkle 8590 LAM!, AHFH!; RSA-POM;
SBBG.

Santa Catalina: RSA-POM; SBBG.

Plantago maritima L. var. californica (Fem.) Pilg.

Santa Rosa: RSA-POM; SBM.

Plantago ovata Forsk.

Santa Rosa: Munz & Crow 1 1 708 LA!; SBBG; SBM.
Anacapa: SBBG; SBM.

San Nicolas: Trask s.n. in Apr. 1897 CAS#955!; Dunkle
8355 LAM!, AHFH!; Kanakoff s.n. Apr. 1 2, 1 940 LAM!;
RSA-POM; SBBG.

Santa Barbara: Dunkle 7430 LAM!, AHFH!; Dunkle 8702
AHFH!; RSA-POM; SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM#28733!;
Trask s.n. in Mar. 1901 LAM#28734!; Fosberg S4777
LAM!; RSA-POM.

San Clemente: Dunkle 7269 LAM!, AHFH!; Dunkle 7270
LAM!, AHFH!; RSA-POM; SBBG.

Guadalupe: Rempel 758-37 LAM!; Palmer 54 CM!; Palm-
er 878 ND-G!.

Platanaceae

Plat anus racemosa Nutt.

Santa Cruz: RSA-POM; SBBG.

Santa Catalina: RSA-POM.

Plumbaginaceae

Armeria maritima (Mill.) Willd. ssp. californica (Boiss.)
G.H.M. Lawr.

Santa Rosa: RSA-POM; SBBG; SBM.

Limonium perezii (Stapf.) F.T. Hubb. ex Bailey
Santa Catalina: RSA-POM; SBBG.

San Clemente: SBBG.

Limonium sinuatum (L.) Mill.

Santa Catalina: RSA-POM.

Polemoniaceae

Allophyllum gilioides (Benth.) A. Grant & V. Grant

Guadalupe: Greene s.n. Apr. 26, 1885 UC!; Palmer 77
MO!, NY!; Greene s.n. Apr. 24, 1885 DS!.

Allophyllum glutinosum (Benth.) A. Grant & V. Grant
Santa Catalina: Fosberg S5413 LAM!, NY!; POM!, SBM!,
US!; Dunkle 2442 AHFH!; Trask s.n. in Mar. 1897 US!;
RSA-POM; SBBG.

Eriastrum filifolium (Nutt.) Woot. & Standi.

Santa Cruz: Greene s.n. in Jul.-Aug. 1886 ND-G#041376!.
Santa Catalina: Trask s.n. in May 1901 LAM!; Thome
36527 RSA!; SBBG.

San Clemente: Raven 17723 RSA!.

Gilia angelensis V. Grant

Santa Cruz: Fosberg 7674 LAM!, POM!; Elmore 440 LAM!;

Raven & Smith 15199 RSA!; CAS; SBBG; SBM.
Anacapa: Blakley 5008 CAS!; SBBG.

Santa Catalina: Moran 697 LAM!, DS!, RSA!; Dunkle 2070
AHFH!; Grant 18438 RSA!; CAS; LA; NY; SBBG; SBM.
San Clemente: SBBG.

74 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Gilia capitata Sims ssp. abrotanifolia (Nutt, ex Greene) V.
Grant

Santa Cruz: Fosberg 7700 LAM!, POM!; SBM.

Santa Catalina: Thorne 36195 RSA!; SBBG.

Gilia clivorum (Jeps.) V. Grant
San Miguel: Munz & Norris 1 1763 POM!; SBBG; SBM.
Santa Rosa: Moran 8 1 8 LAM!, RSA!; Munz & Crow 1 1 696
POM!; Raven, Blakley & Ornduff 14907 RSA!; CAS;
LA; SBBG; SBM.

Santa Cruz: Fosberg 7522 LAM!; Breedlove 2818 RSA!;

Raven & Smith 15128 RSA!; CAS; SBBG; SBM.
Anacapa: Moran 722 LAM!; Blakley 4944 CAS!; Blakley
5013 CAS!; SBBG; SBM.

Santa Catalina: Fosberg S4538 LAM!.

Gilia nevinii Gray
Santa Rosa: SBBG.

Santa Cruz: Brandegee s.n. in Apr. 1888 CAS!; SBBG.
Anacapa: Blakley 4950 CAS!; Blakley 4975 CAS!; SBBG.
San Nicolas: Trask 53 MO!; Trask 52 NY!; Trask s.n. in
Apr. 1897 CAS!.

Santa Barbara: Dunkle 7429 LAM!; Thome 37503 RSA!;
Moran 830 DS!; SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!, NY!; Fos-
berg S4496 LAM!; Trask s.n. in Apr. 1900 CAS#473!;
RSA-POM; SBBG.

San Clemente: Dunkle 7320 LAM!; Trask 248 NY!, US!;

Raven 17213 RSA!; DS; LA; SBBG.

Guadalupe: Palmer 78 NY!; Greene s.n. Apr. 25, 1885
CAS#36173!; Moran 17289 RSA!; GH.

Gilia tenuiflora Benth. ssp. hoffmannii (Eastw.) A. Grant &
V. Grant

Santa Rosa: Moran 819 LAM!, RSA!; Moran 793 LAM!,
RSA!; Munz & Hoffmann 11730 POM!; SBM.
Linanthus androsaceus (Benth.) Greene ssp. luteus (Benth.)
Mason

Santa Cruz: SBBG.

Linanthus bicolor (Nutt.) Greene ssp. bicolor
Santa Rosa: Thome et al. 48860 RSA!.

Santa Catalina: Dunkle 1795 AHFH!, POM!; Trask s.n.

in Mar. 1901 LAM!, NY!; Dunkle 2106 AHFH!.

San Clemente: Dunkle 7262 LAM!, AHFH!; Munz 6622
POM!; Raven 17226 RSA!; SBBG.

Linanthus dianthiflorus (Benth.) Greene ssp. dianthiflorus
Santa Cruz: Blakley 3310 RSA!; Wolf 2790 RSA!; SBBG;
SBM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Moran 695
LAM!; Dunkle 1740 AHFH!, POM!; SBBG.

Linanthus pygmaeus (Brand) J.T. Howell ssp. pygmaeus
San Clemente: Thome 36077 RSA!; Raven 17692 RSA!;
SBBG.

Guadalupe: Moran 6644 RSA!; Moran 17388 RSA!;
Carlquist 464 RSA!.

Navarretia atractyloides (Benth.) H. & A.

Santa Rosa: Dunkle 8507 LAM!; Epling & Erickson s.n.

Aug. 8, 1937 LA!; SBM.

Santa Cruz: Dunkle 8559 AHFH!; SBBG; SBM.

Santa Catalina: Davidson, A. s.n. Jun. 26, 1891 LAM!;
Pendleton 1392 POM!; Nuttall s.n. UC!; SBBG.

San Clemente: House & Grumbles s.n. Aug. 5-13, 1930
USC!.

Navarretia hamata Greene var. foliacea (Greene) Thorne
Santa Catalina: Dunkle 1920 AHFH!, POM!; Fosberg
S4674 LAM!; Wolf 3547 LAM!, RSA!; SBM.
Navarretia hamata Greene var. hamata
Santa Cruz: RSA-POM; SBBG.

Santa Catalina: RSA-POM; SBBG.

San Clemente: Raven 17724 RSA!; Blakley 3311 RSA!;
Clokey 5032 POM!; SBBG.

Polygalaceae

Polygala californica Nutt.

Santa Cruz: Hoffmann 188 POM!; Raven & Smith 15224
RSA!; SBBG.

Polygonaceae

Chorizanthe coriaceae Goodm.

Santa Rosa: Munz & Hoffman 1 1734 POM!; SBM.

Santa Cruz: SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!, NY!; Thome
& Everett 34600 RSA!.

Chorizanthe staticoides Benth. ssp. staticoides
Santa Cruz: Clokey 4907 LAM!, POM!; Balls & Blakley
23742 RSA!.

Santa Catalina: Fosberg S4917 LAM!, POM!; Trask s.n.
in May 1901 NY!; Trask s.n. in Jun. 1897 US!.
Chorizanthe wheeleri Wats.

Santa Rosa: Munz & Crow 1 1679 POM!; Raven, Blakley
& Ornduff 15010 RSA!; SBM.

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 CAS!, LA!, POM!;
Munz & Norris 11847 POM!; Raven & Smith 15182
RSA!.

Eriogonum arborescens Greene

Santa Rosa: Dunkle 8449 LAM!, AHFH!; Dunn, N. s.n.

May 15, 1932 LA!; Thome et al. 49005 RSA!; SBBG.
Santa Cruz: Yates s.n. in 1893 LAM!; Hutchinson s.n.
LAM!; Clokey 4910 LAM!, POM!; LA; RSA-POM;
SBBG; SBM.

Anacapa: Dunkle 7618 LAM!, AHFH!, RSA!; Howell 3791
USC!; Bond 321 SBM!.

Eriogonum cinereum Benth.

Santa Rosa: RSA-POM.

Eriogonum fasciculatum Benth. ssp .fasciculatum
Santa Catalina: Thome, Propst & Haefs 45110 RSA!.
Eriogonum giganteum Wats. ssp. compaction (Dunkle) Munz
Santa Barbara: Dunkle 8103 LAM!, AHFH!; Bryan, Dr.
& Mrs. s.n. LAM!; Elmore 306 AHFH!; RSA-POM;
SBBG; SBM.

Eriogonum giganteum Wats. ssp. formosum (K. Bdg.) Raven
San Clemente: Moran 6843 LA!; Brandegee s.n. Aug. 25,
1894 DS#88608!, DS#88624!; RSA-POM; SBBG.
Eriogonum giganteum Wats. ssp. giganteum

Santa Cruz: RSA-POM; SBBG (probably introduced here).
Santa Catalina: Davidson, A. s.n. Jun. 26, 1891 LAM!;
Grant s.n. Sep. 1, 1906 LAM!; Fosberg S5358 LAM!,
SBM!; RSA-POM; SBBG.

Eriogonum grande Greene var. dunklei Reveal
San Miguel: Dunkle 8369 LAM!, AHFH!, DS!; Elmore
325 AHFH!; Elmore 334 AHFH!.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 75

Eriogonum grande Greene ssp. grande
Santa Rosa: Hoffmann 133 SBM!.

Santa Cruz: Clokey 4908 LAM!; Greene s.n. in Jul.-Aug.
1886 CAS#336!; Dunkle 8594 LAM!, AHFH!; RSA-
POM.

Anacapa: Hoffmann 93 SBM; Dunkle 7612 LAM!, AHFH!;

Elmore 232 AHFH!; RSA-POM; SBBG; SBM.

San Nicolas: SBBG.

Santa Catalina: Fosberg 7150 LAM!; Dunkle 2030 LAM!;

Davidson, A. s.n. LAM#20543!; RSA-POM.

San Clemente: Dunkle 7290 LAM!, AHFH!; House &
Grumbles s.n. Aug. 5-13, 1930 USC!; Johnstone s.n.
Sep. 5, 1926 USC!; RSA-POM; SBBG.

Eriogonum grande Greene ssp. rubescens (Greene) Munz
San Miguel: Greene s.n. in Sep. 1886 CAS#88643!; RSA-
POM; SBBG; SBM.

Santa Rosa: Dunkle 8442 LAM!, AHFH!; Dunn, N. s.n.
May 24, 1931 LAM!; Hoffmann 92 SBM!; LA; RSA-
POM; SBBG.

Santa Cruz: Hoffmann s.n. SBM#629!; RSA-POM; SBBG.
Anacapa: Hoffmann 85 SBM!; Hoffmann 86 SBM!; Yates
s.n. SBM#17301!.

Eriogonum grande Greene ssp. timorum (Reveal) Munz
San Nicolas: Moran 3168 LA!; Foreman 213 LA!; Raven
& Thompson 20744 LA!; RSA-POM.

Eriogonum zapatoense Moran

Guadalupe: Moran 18170 LAM!, CAS!, RSA!; Rempel
759-37 LAM!.

Polygonum arenastrum Bor.

Santa Rosa: Hoffmann s.n. Apr. 14, 1929 SBM#6286!;

Hoffmann s.n. Jun. 12, 1930 SBM#8754!.

Santa Cruz: Hoffmann s.n. Jun. 14, 1930 SBM#5111!;

Hoffmann s.n. Sep. 20, 1930 SBM# 10306!.

Santa Catalina: Fosberg S5410 LAM!; SBBG.

San Clemente: SBBG.

Polygonum argyrocoleon Steud. ex Kunze
Santa Cruz: Hobbs 78-1 16 LA!.

Santa Catalina: Fosberg S4461 LAM!; Fosberg S4565
LAM!; Spalding s.n. Sep. 19, 1925 LAM!; RSA-POM;
USC.

San Clemente: House & Grumbles s.n. Aug. 5-13, 1930
LAM!.

Polygonum aviculare L.

Santa Rosa: Dunkle 8453 LAM!; Elmore 213 AHFH!;

Hoffmann s.n. Jul. 7, 1930 SBM#10398!.

Santa Cruz: Dunkle 8628 LAM!, AHFH!; Hoffmann s.n.

SBM#10306!; RSA-POM.

Santa Catalina: RSA-POM.

San Clemente: RSA-POM.

Pterostegia drymarioides F. & M.

San Miguel: Hoffmann s.n. SBM#9225!; RSA-POM.
Santa Rosa: Dunn, N. s.n. May 24, 1931 LA!; Hoffmann
s.n. SBM#141 1!; Hartwell s.n. SBM# 14372!; RSA-POM.
Santa Cruz: Clokey 4904 LAM!; Dunkle 8619 LAM!; Fos-
berg 7620 LAM!, LA!; RSA-POM.

Anacapa: Hoffmann s.n. SBM#5572!; SBBG.

Santa Barbara: Dunkle 7414 LAM!; Bond s.n. SBM# 15095!;
RSA-POM.

Santa Catalina: Dunkle 2094 AHFH!; Fosberg S4452 (in
part) LAM!; Dunkle 1825 AHFH!; RSA-POM.

San Clemente: Moran 576 LAM!; Dunkle 7207 AHFH!,
LAM!.

Guadalupe: Palmer 843 ND-G!; Norris s.n. May 1, 1951
LA!.

Rumex angiocarpus Murbeck

Santa Cruz: Hoffmann s.n. Mar. 20, 1932 SBM#11767!;
Daily 615 SCIR!.

Rumex conglomeratus Murr.

Santa Rosa: RSA-POM.

Santa Cruz: Daily 542 SCIR!.

Santa Catalina: Fosberg S5369 LAM!; RSA-POM.
Rumex crispus L.

San Miguel: Dunkle 8394 LAM!, AHFH!; Hoffmann s.n.
SBM#5001!; RSA-POM.

Santa Rosa: Hoffmann s.n. SBM#6257!; RSA-POM.
Santa Cruz: Dunkle 8586 LAM!; Hoffmann s.n.

SBM#6642!; RSA-POM.

Anacapa: Hoffmann s.n. SBM# 1788!; SBBG.

San Nicolas: RSA-POM.

Santa Catalina: Dunkle 1919 AHFH!; Trask s.n. in Mar.

1901 LAM!; Fosberg S4965 LAM!; RSA-POM.

San Clemente: Elmore 416 LAM!, AHFH!; RSA-POM.
Rumex fueginus Phil.

San Miguel: Norris s.n. SBM#9265!; RSA-POM.

Rumex pulcher L.

Santa Cruz: Hoffmann s.n. Apr. 11, 1931 LAM!; Hoffmann
s.n. SBM#7226!; SBBG.

Rumex salicifolius Weinm.

San Miguel: SBBG.

Santa Rosa: Dunkle 8475 LAM!; Dunkle 8522 LAM!,
AHFH!; Hoffmann s.n. SBM# 10063!.

Santa Cruz: Clokey 4903 LAM!; Hoffmann s.n. SBM#52 1 3!;
RSA-POM.

San Nicolas: Dunkle 8331 LAM!; Raven & Thompson
20729 LA!; RSA-POM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Dunkle
1986 AHFH!; Fosberg S4708 LAM!, SBM!; RSA-POM.
San Clemente: RSA-POM.

Portulacaceae
Calandrinia breweri Wats.

Santa Rosa: SBM.

Santa Cruz: Fosberg 7461 LAM!; RSA-POM; SBM.
Calandrinia ciliata (R. & P.) DC. var. menziesii (Hook.)
Macbr.

San Miguel: SBBG.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: RSA-POM; SBM.

Anacapa: SBBG; SBM.

Santa Barbara: SBBG.

Santa Catalina: RSA-POM.

San Clemente: RSA-POM.

Guadalupe: Palmer 14 CM!; Norris s.n. May 1, 1951
LA#95561!; Moran 6635 RSA!.

Calandrinia maritima Nutt.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

76 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Anacapa: SBBG.

Santa Barbara: Dunkle 7406 LAM!; Dunkle 7432 LAM!;
RSA-POM; SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; RSA-POM.
San Clemente: RSA-POM; SBBG.

Guadalupe: Carlquist 467 RSA!.

Clavtonia perfoliata Donn var. parviflora (Dougl. ex Hook.)
Torr.

Santa Cruz: RSA-POM; SBBG.

Claytonia perfoliata Donn var. perfoliata
San Miguel: RSA-POM; SBM.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: Fosberg 7586 LAM!, LA!; Clokey 4927 LAM!;

Clokey 4925 LAM!; RSA-POM; SBBG; SBM.
Anacapa: SBBG; SBM.

San Nicolas: RSA-POM.

Santa Barbara: Dunkle 7457 LAM!, AHFH!; Dunkle 7407
LAM!, AHFH!; SBBG; SBM.

Santa Catalina: Fosberg S4486 LAM!; Fosberg S4435
LAM!; Dunkle 1797 AHFH!; RSA-POM.

San Clemente: Elmore 418 AHFH!; Dunkle 7224 LAM!;
RSA-POM.

Guadalupe: Palmer 846 ND-G!; Norris s.n. May 1, 1951
LA#95563!.

Montia fontana L. ssp. amporitana Sennen
Santa Cruz: SBM.

Portulaca oleracea L.

Santa Cruz: Hoffman s.n. in 1930 SBM#4187!.

Santa Catalina: RSA-POM.

Talinum guadalupense Dudley in D.S. Jordan
Guadalupe: Rempel 759-37 LAM!; Thobum, Greene &
Wing s.n. in Jul. 1897 DS#140708!; Moran 2635 RSA!.
Primulaceae
Anagallis arvensis L.

Santa Rosa: Elmore 208 LAM!; Dunkle 8454 LAM!; RSA-
POM; SBBG; SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Anacapa: SBBG.

Santa Catalina: Fosberg S4433 LAM!; Nuttall 5 NY!; Trask
s.n. in 1898 US!; AHFH; USC.

Anagallis minima (L.) Krause
Santa Rosa: SBM.

Dodecatheon clevelandii Greene ssp. insularis H.J. Thomp-
son

Santa Rosa: SBBG; SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Anacapa: Moran 713 LAM!; SBBG; SBM.

Santa Catalina: Moran 604 LAM!; Dunkle 2054 LAM!,
AHFH!; Fosberg S4361 LAM!; SBBG; SBM.

San Clemente: Moran 571 LAM!; SBBG; SBM.
Guadalupe: Palmer 55 CM!; Moran 6464 SD!; Moran
21166 SD!.

Samolus parviflorus Raf.

Santa Cruz: Hoffmann s.n. Sep. 10, 1931 LAM!; SBM.
Ranunculaceae

Clematis lasiantha Nutt, in T. & G.

Santa Cruz: Hoffmann 218 POM!; Wolf 2755 RSA!, US!;
Raven & Smith 15275a RSA!; CAS; SBBG; SBM.

Clematis ligusticifolia Nutt, in T. & G.

Santa Rosa: Dunkle 8431 LAM!, AHFH!, RSA!; SBBG;
SBM.

Santa Cruz: Clokey 4937 LAM!, POM!; Dunkle 8649 LAM!,
AHFH!, RSA!; Williams 99 POM!; CAS; SBBG; SBM.
Santa Catalina: Fosberg S5367 LAM!; Dunkle 1990
AHFH!, POM!; Dunkle 1722 AHFH!; SBBG; SBM.
Delphinium kinkiense Munz

San Clemente: Dunkle 7322 LAM!, RSA!; Beauchamp 290
RSA!.

Delphinium parryi Gray ssp. parryi
San Miguel: SBM.

Santa Rosa: Hoffmann 129 POM!; Youngberg s.n. May
29, 1938 POM!; Dunn, A.N. & A.M. Martin s.n. May
15, 1932 LA!; SBBG; SBM.

Santa Cruz: Clokey 4936 LAM!; Hoffmann Apr. 10, 1931
LAM!; Breedlove 2861 RSA!; SBBG; SBM.

Anacapa: Moran 714 LAM!, RSA!; SBBG; SBM.

Santa Catalina: Moxley 593 LAM!; Smith, R.J. s.n. May
24, 1934 LAM!; Moran 667 LAM!; LA; RSA-POM;
SBBG.

San Clemente: Raven 17820 RSA!.

Delphinium variegatum T. & G. ssp. thornei Munz

San Clemente: Raven 17700 LA!; Thorne 42801 RSA!;
SBBG.

Myosurus minimus L. var. filiformis Greene

Guadalupe: Greene s.n. Apr. 19, 1885 ND-G#01 8340!.
Ranunculus californicus Benth. var. californicus

San Miguel: Schuyler 29 LAM!; Youngberg s.n. May 28,
1938 POM!; SBBG; SBM.

Santa Rosa: Moran 817 LAM!; Dunkle 8488 AHFH!;

Thome et al. 48995 RSA!; LA; SBBG; SBM.

Santa Cruz: Moran 751 LAM!; Fosberg 7690 LAM!; Clo-
key 5184 NY!, RSA!; LA; SBBG; SBM.

Ranunculus californicus Benth. var. cuneatus Greene
San Miguel: Hoffmann s.n. Apr. 19, 1932 POM!; Munz &
Norris 11880 POM!; SBM.

Santa Cruz: Wolf 2798 RSA!; SBM.

Ranunculus hebecarpus H. & A.

Santa Catalina: Fosberg S4348 LAM!, POM!; Detmers s.n.

Apr. 13, 1929 USC!; SBM.

Guadalupe: Moran 6610 RSA!; Carlquist 443 RSA!.
Resedaceae

Oligomeris linifolia (Vah.) Macbr.

San Miguel: SBBG; SBM.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: SBM.

Anacapa: SBBG.

San Nicolas: Trask s.n. in Apr. 1900 LAM!; Dunkle 8318
LAM!, AHFH!; Raven & Thompson 20741 LA!; RSA-
POM; SBBG.

Santa Barbara: Bryan, Dr. & Mrs. s.n. Jul. 14, 1922 LAM!;
RSA-POM; SBBG.

Santa Catalina: Davidson, A. s.n. Jun. 26, 189- LAM!;

RSA-POM; SBBG.

San Clemente: RSA-POM; SBBG.

Guadalupe: Moran 5955 LA!, RSA!; Carlquist 471 RSA!;
Palmer 10 CM!.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 77

Reseda odorata L.

Santa Catalina: RSA-POM.

Rhamnaceae

Ceanothus arboreus Greene

Santa Rosa: Munz & Crow 11611 LA!; RSA-POM; SBM.
Santa Cruz: Fosberg 7577 LAM!; Dunkle 8658 LAM!;

Moran 747 LAM!; LA; CAS; RSA-POM; SBBG; SBM.
Santa Catalina: Trask s.n. in Mar. 1900 LAM!; Davidson,
A. s.n. LAM!; Grant & Wheeler s.n. Apr. 21-26, 1904
LAM!; LA; RSA-POM; SBBG; SBM.

Ceanothus crassifolius Torr.

Guadalupe: Franceschi s.n. UC!; Palmer 22 MO!, NY!.
Ceanothus megacarpus Nutt. ssp. insu/aris (Eastw.) Raven
Santa Rosa: SBBG.

Santa Cruz: Dunkle 8650 LAM!; Fosberg 7559 LAM!;

Clokey 5001 DS!, NY!; LA; RSA-POM; SBBG; SBM.
Anacapa: Dunkle 7665 LAM!, AHFH!; RSA-POM; SBBG;
SBM.

Santa Catalina: Fosberg S4920 LAM!; Trask s.n. in Mar.

1 90 1 NY!; Millspaugh 4590 F!; CAS; RSA-POM; SBBG.
San Clemente; Trask 198 NY!; Blakley 5197 DS!; RSA-
POM; SBBG.

Ceanothus megacarpus Nutt. ssp. megacarpus
Santa Cruz: SBBG.

Santa Catalina: Dunkle 2038 LAM!; Dunkle 2059 LAM!;
Lister & Powell s.n. Apr. 5, 1928 USC!; CAS; DS; RSA-
POM; SBBG.

San Clemente: Murbarger 165 UC!.

Rhamnus californica Esch. ssp. californica
Santa Cruz: Hoffmann 669 SBM!; RSA-POM.

Rhamnus pirifolia Greene
Santa Rosa: RSA-POM; SBM.

Santa Cruz: Hoffmann s.n. in Jun. 1930 LAM!; Eastwood
6434 GH!; Clokey 4997 DS!, NY!; CAS; RSA-POM;
SBBG; SBM.

Santa Catalina: Fosberg S4238 LAM!, NY!; Grant s.n.
LAM!; Sargent s.n. Sep. 16, 1874 GH!; AHFH; DS; LA;
RSA-POM; SBBG; SBM.

San Clemente: Trask 109 NY!; Moran, Beauchamp &
Oberbauer 22682 CAS!; RSA-POM; SBBG.

Guadalupe: Moran 13803 LAM!; Palmer 21 NY!; Moran
12036 RSA!.

Rosaceae

Adenostoma fasciculatum H. & A. var. fasciculatum

Santa Rosa: Dunkle 8496 LAM!; Dunn, N. s.n. May 24,
1931 LA!; RSA-POM; SBBG; SBM.

Santa Cruz: Dunkle 8609 LAM!, AHFH!; Dunkle 8657
LAM!, AHFH!; Fausett 20 LA!; RSA-POM; SBBG; SBM.
Santa Catalina: deForest s.n. May 19, 1934 LAM!; Fosberg
S4775 LAM!; Fosberg S4562 LAM!; RSA-POM; SBBG.
San Clemente: Dunkle 7347 LAM!, AHFH!; RSA-POM;
SBBG.

Alchemil/a occidentalis Nutt.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7590 LAM!; RSA-POM; SBBG; SBM.
Santa Catalina: Fosberg S4592 LAM!; Fosberg S4341
LAM!; RSA-POM; SBBG.

San Clemente: RSA-POM.

Guadalupe: Palmer 29 CM!; Wiggins & Ernst 34 DS!; Mo-
ran 2913 DS!.

Cercocarpus betuloides Nutt, ex T. & G. ssp. betuloides
Santa Cruz: Fosberg 7514 LAM!, LA!; Clokey 4957 LA!;
SBM.

Santa Catalina: Fosberg S4798 LAM!; Dunkle 1711
AHFH!.

Cercocarpus betuloides Nutt, ex T. & G. ssp. blancheae (C.
K. Schneid.) Thome
Santa Rosa: RSA-POM.

Santa Cruz: Hoffmann 185 LAM!; Yates s.n. in Aug. 1893
LAM!; Fosberg 7627 LAM!, LA!; RSA-POM; SBBG;
SBM.

Santa Catalina: Blakley 5478 LAM!; Moran 703 LAM!;
Dunkle 2057 LAM!, AHFH!; LA; RSA-POM; SBBG.
Cercocarpus traskiae Eastw.

Santa Catalina: Trask s.n. in Mar. 1897 CAS# 141!; Trask
s.n. in Apr. 1903 LAM!; Moran 678 LAM!; LA; RSA-
POM; SBBG.

Heteromeles arbutifolia (Ait.) M. Roem.

San Miguel: SBBG.

Santa Rosa: Dunkle 8486 LAM!, AHFH!; Elmore 210
AHFH!; Dunn, N. s.n. May 24, 1931 LA!; RSA-POM;
SBBG; SBM.

Santa Cruz: Dunkle 8535 LAM!, AHFH!; Elmore 269
AHFH!; Eastwood 6380 MO!; LA; RSA-POM; SBBG;
SBM.

Anacapa: Dunkle 7630 LAM!, AHFH!; Johnstone s.n.

USC!; RSA-POM; SBBG; SBM.

Santa Catalina: Templeton 11397 LAM!; Fosberg S4445
LAM!; Doushel s.n. Jul. 23, 1915 MO!; LA; RSA-POM;
SBBG; SBM.

San Clemente: Dunkle 7337 AHFH!; Elmore 421 AHFH!;

DeBuhr & Wallace 701 LAM!; RSA-POM; SBBG; USC.
Guadalupe: Newcomb 184 DS!; Wiggins & Ernst 202 DS!.

Holodiscus discolor (Pursh) Maxim, var. discolor
Santa Cruz: Hoffmann 9 POM!; SBBG.

Santa Catalina: Fosberg S5436 LAM!, POM!; Thome
36905 RSA!; SBBG.

Lyonothamnus floribundus Gray ssp. asplenifolius (Greene)
Raven

Santa Rosa: Munz & Voss 1 1569 LA!; RSA-POM; SBBG;
SBM.

Santa Cruz: Yates s.n. in Aug. 1893 LAM!; Fosberg 7655
LAM!; Dunkle 8611 LAM!; LA; RSA-POM; SBBG;
SBM.

San Clemente; Dunkle 7316 LAM!, AHFH!; DeBuhr &
Wallace 712 LAM!; RSA-POM; SBBG.

Lyonothamnus floribundus Gray ssp. floribundus

Santa Catalina: Trask s.n. in Jun. 1897 LAM!; Fosberg
S4686 LAM!; Moran 652 LAM!; LA; RSA-POM; SBBG.
Potenti/la egedii Wormsk. var. grandis (Rydb.) J.T. Howell
San Miguel: RSA-POM.

Santa Cruz: RSA-POM.

Potentilla glandulosa Lindl. ssp. glandulosa
Santa Catalina: RSA-POM.

78 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Prunus lyonii (Eastw.) Sarg.

Santa Rosa: Dunkle 8510 LAM!, AHFH!; Dunn 1 326 LA!;
Epling & Erickson s.n. Aug. 8, 1937 LA!; RSA-POM;
SBBG; SBM.

Santa Cruz: Moran 754 LAM!; Dunkle 8556 LAM!,
AHFH!; Elmore 271 AHFH!; LA; RSA-POM; SBBG;
SBM.

Anacapa: Dunkle 763 1 LAM!, AHFH!; Moran 732 LAM!;
RSA-POM; SBBG; SBM.

Santa Catalina: Davidson, A. s.n. LAM!; Fosberg S4488
LAM!; Dunkle 1756 AHFH!; LA; RSA-POM; SBBG;
SBM.

San Clemente: DeBuhr & Wallace 697 LAM!; Dunkle 7343
LAM!, AHFH!; Dunkle 7243 AHFH!; RSA-POM;
SBBG.

Prunus persica (L.) Batsch
Santa Catalina: RSA-POM.

Rosa californica Cham. & Schlecht.

Santa Rosa: Dunkle 8434 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

Santa Cruz: Dunkle 8647 LAM!, AHFH!; Elmore 447
AHFH!; Hoffmann s.n. Jul. 1, 1930 LAM!; RSA-POM;
SBBG; SBM.

Santa Catalina: Davidson, A. s.n. in Jun. 1892 LAM!;
Fosberg S4743 LAM!; Dunkle 1946 AHFH!; RSA-POM.
Rubus procerus P.J. Muell.

Santa Catalina: RSA-POM.

Rubus ursinus Cham. & Schlecht.

San Miguel: SBM.

Santa Rosa: Dunkle 8448 AHFH!; RSA-POM; SBBG;
SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Fosberg S4740 LAM!; Davidson, A. s.n.
in Jun. 1891 LAM!; Moran 685 LAM!; RSA-POM;
SBBG.

Rubiaceae

Galium angulosum Gray

Guadalupe: Palmer 36 MO!, CM!; Moran 18149 MO!;
Carlquist 453 RSA!.

Galium angustifolium Nutt, ex T. & G. ssp. angustifolium
Santa Catalina: Fosberg S4870 LAM!; Trask s.n. in May
1900 LAM!; Hasse 4135 NY!; LA; RSA-POM; SBBG;
SBM.

Galium angustifolium Nutt, ex T. & G. ssp .foliosum (Hilend
& Howell) Dempst. & Steb.

Santa Rosa: Dunkle 8447 LAM!, AHFH!; RSA-POM; SBM.
Santa Cruz: Hoffmann s.n. Sep. 10, 1931 LAM!; Howell
6196 LA!; Greene s.n. in Jul. -Aug. 1886 NY!; RSA-
POM; SBBG; SBM.

Anacapa: Johnstone s.n. Jun. 25, 1932 LAM!; Hoffmann
s.n. Jun. 16, 1930 LAM!; Dunkle 7615 LAM!, AHFH!;
LA; RSA-POM; SBBG; SBM.

Galium aparine L.

San Miguel: RSA-POM; SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7668 LAM!, LA!; RSA-POM; SBM.
Anacapa: SBBG; SBM.

San Nicolas: RSA-POM; SBBG.

Santa Barbara: Dunkle 7417 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

Santa Catalina: Fosberg S4613 LAM!; Fosberg S4311
LAM!; Dunkle 2087 AHFH!; RSA-POM; SBBG; SBM.
San Clemente: Dunkle 7220 LAM!, AHFH!; Moran 581
LAM!; Dunkle 7311 AHFH!; RSA-POM; SBBG.
Guadalupe: Palmer 35 MO!, NY!; Palmer 850 ND-G!.
Galium buxifolium Greene

San Miguel: Hoffmann s.n. Jun. 11, 1930 LAM!, CAS!;

RSA-POM; SBBG; SBM.

Santa Rosa: Moran 807 RSA!.

Santa Cruz: Fosberg 7666 LAM!, LA!; Abrams & Wiggins
74 NY!; Yates s.n. SBM#2 1111!; CAS; RSA-POM;
SBBG.

Galium californicum H. & A. ssp. flaccidum (Greene) Dempst.
Santa Cruz: Dunkle 8607 LAM!, AHFH!; Clokey 5145
LA!; Hoffmann 252 LA!; RSA-POM; SBBG; SBM.
Galium californicum H. & A. ssp. miguelense (Greene) Demp.
& Steb.

San Miguel: SBBG; SBM.

Santa Rosa: Smith 8240 NY!; RSA-POM; SBBG; SBM.
Galium catalinense Gray ssp. acrispum Dempst.

San Clemente: Trask 191 NY!; Trask 1 92 NY!; RSA-POM.
Galium catalinense Gray ssp. catalinense

Santa Catalina: Davidson, A. s.n. Jun. 20, 1891 LAM!;
Grant & Wheeler s.n. Apr. 21-26, 1904 LAM!, NY!;
Beauchamp 376 LAM!; RSA-POM; SBBG.

Galium nuttallii Gray ssp. insulare Ferris
Santa Rosa: Moran 807 LAM!, CAS!, DS!, NY!; Dunn, N.

s.n. May 24, 1931 LA!; RSA-POM; SBBG; SBM.

Santa Cruz: Clokey 5148 LA!, NY!; Clokey 5076 NY!;

Howell 6362 NY!; RSA-POM; CAS; DS; SBBG; SBM.
Santa Catalina: Moran 606 LAM!, NY!; Moran 700 LAM!,
DS!, NY!; Fosberg S4688 LAM!; RSA-POM; SBBG.
Galium porrigens Dempst. var. porrigens
Santa Rosa: Blakley 3055 SBBG!; Blakley 3147 SBBG!.
Santa Cruz: Ferren 1879 SCIR!; RSA-POM.

Rutaceae

Ruta chalepensis L.

Santa Cruz: Daily 623 SCIR!.

Santa Catalina: Millspaugh 4647 F!.

Guadalupe: Moran 12019 SD!.

Salicaceae

Populus fremontii Wats, ssp . fremontii
Santa Cruz: SBM.

Santa Catalina: Fosberg S4853 LAM!.

Populus trichocarpa T. & G.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: Fosberg 7560 LAM!; Hoffmann s.n. Sep. 8,
1931 LAM!; SBBG; SBM.

Santa Catalina: Fosberg S4830 LAM!; Fosberg S4704
LAM!; Dunkle 1804 AHFH!; SBM.

Populux x parryi Sarg.

San Nicolas: Raven & Thompson 20728 DS!.

Santa Catalina: RSA-POM.

Salix hindsiana Benth. var. hindsiana
Santa Cruz: RSA-POM.

Santa Catalina: Nuttall 343 F!.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 79

Salix laevigata Bebb. var. laevigata
Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!, NY!; Da-
vidson, A. s.n. LAM#9468!; Dunkle 1 730 AHFH!; RSA-
POM; SBBG.

Salix iasiandra Benth. var. lasiandra
Santa Cruz: RSA-POM; SBM.

Salix iasiolepis Benth. var. lasiolepis
San Miguel: SBBG.

Santa Rosa: Dunkle 8516 LAM!, AHFH!, NY!; RSA-POM;
SBBG; SBM.

Santa Cruz: Dunkle 8636 LAM!, AHFH!; Fosberg 7705
LAM!; Fosberg 7629 LAM!; RSA-POM; SBBG; SBM.
San Nicolas: Blakley 4090 SBBG!; Blakley 4163 SBBG!;

Raven & Thompson 20733 DS!; RSA-POM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Fosberg
S4553 LAM!; Dunkle 2075 AHFH!; LA; RSA-POM;
SBBG.

Saururaceae

Anemopsis californica (Nutt.) H. & A.

Santa Cruz: Clokey 4902 LAM!, NY!; Rowntree s.n. Jun.
1 5, 1 930 SBM!; Abrams & Wiggins 1 1 6 CAS!, DS!; RSA-
POM; SBBG.

San Nicolas: Trask 98 MO!, NY!, US!.

Santa Catalina: Dunkle 1926 AHFH!; Wolf 3587 US!,
DS!; RSA-POM; SBBG.

San Clemente: (Raven 1963: Murbarger 143 at UC).
Saxifragaceae
Heuchera maxima Greene

Santa Rosa: Dunkle 8524 LAM!; Epling & Erickson s.n.

Aug. 8, 1937 LA!; RSA-POM; SBBG; SBM.

Santa Cruz: Clokey 4952 LAM!; Dunkle 8618 LAM!;
Greene s.n. in Jul.-Aug. 1886 CAS#23479!; RSA-POM;
SBBG; SBM.

Anacapa: Dunkle 7653 LAM!; Moran 717 LAM!; RSA-
POM; SBBG; SBM.

Jepsonia malvaefolia (Greene) Small
Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7607 LAM!; Kellogg & Harford s.n.

in 1874 CAS#631!; RSA-POM; SBM.

San Nicolas: Howell 8210 CAS!.

Santa Catalina: Dunkle 2019 AHFH!; RSA-POM; SBBG.
San Clemente: Dunkle 7258 LAM!, AHFH!; Dunkle 7356
AHFH!; RSA-POM.

Guadalupe: Palmer s.n. in 1885 GH!; Moran 2917 DS!.
Lithophragma affine Gray ssp. mixtion R.L. Taylor

Santa Catalina: Dunkle 2124 LAM!, AHFH!; Moran 682
LAM!; Fosberg S4581 LAM!; RSA-POM.
Lithophragma cymbalaria T. & G.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Moran 746 LAM!; Fosberg 7624 LAM!, LA!;
Elmore s.n. Apr. 18, 1936 AHFH!; RSA-POM; SBBG;
SBM.

Lithophragma maximum Bacig.

San Clemente: Murbarger 1 18 UC!.

Ribes malvaceum Sm. in Rees var. malvaceum
Santa Rosa: SBM.

Santa Cruz: Hoffmann 1 90 LAM!; RSA-POM; SBBG; SBM.

Anacapa: Dunkle 7670 LAM!; Dunkle 7669 AHFH!; SBBG;
SBM.

San Clemente: Dunkle 7338 LAM!; RSA-POM; SBBG.
Ribes menziesii Pursh var. menziesii
Santa Cruz: RSA-POM; SBM.

Ribes menziesii Pursh var. thacherianum Jeps.

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 LAM!; Elmore
s.n. Apr. 18, 1936 AHFH!; RSA-POM; SBBG.

Ribes sanguineum Pursh

Guadalupe: Palmer s.n. in 1875 GH!

Ribes viburnifolium Gray

Santa Catalina: Merritt s.n. in Apr. 1894 LAM!; Fosberg
S4774 LAM!; Moran 615 LAM!; RSA-POM; SBBG.
Saxifraga californica Greene
Santa Rosa: SBBG.

Santa Cruz: Moran 761 LAM!; Fosberg 7585 LAM!; Plun-
kett s.n. Apr. 18, 1936 LA!; RSA-POM; SBBG; SBM.
Scrophulariaceae
Antirrhinum kelloggii Greene

Santa Cruz: Fosberg 7678 LAM!; RSA-POM; SBBG; SBM.
Santa Catalina: Trask s.n. in Mar. 1 897 LAM!; RSA-POM.
Antirrhinum kingii Wats. var. watsoni (Vasey & Rose) Munz
Guadalupe: Moran 17408 SD!; Moran 17345 SD!.
Antirrhinum multiflorum Penn.

Santa Cruz: SBM.

Antirrhinum nuttallianum Benth. in DC.

San Miguel: SBM.

Santa Rosa: Epling & Erickson s.n. Aug. 8, 1937 LA!; RSA-
POM; SBBG; SBM.

Santa Cruz: Dunkle 8605 LAM!, AHFH!; Yates s.n. in
Aug. 1893 LAM!; Eastwood 6398 LA!; RSA-POM;
SBBG; SBM.

Anacapa: SBBG.

Santa Catalina: Trask s.n. in Mar. 1900 LAM!; Dunkle
1 903 AHFH!; Fosberg S539 1 LAM!; RSA-POM; SBBG.
San Clemente: RSA-POM; SBBG.

Guadalupe: Moran 6593 RSA!; Carlquist 451 RSA!.
Castilleja affinis H. & A. var. affinis
Santa Rosa: Dunkle 8445 LAM!; Dunn, N. s.n. May 24,
1931 LA!; RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7698 LAM!; Hoffmann s.n. Apr. 12,
1931 LAM!; Fosberg 7634 LAM!, LA!; RSA-POM;
SBBG; SBM.

Anacapa: Dunkle 7661 LAM!, AHFH!, POM!; Moran 742
LAM!, NY!; Elmore 230 AHFH!; LA; SBBG; SBM.
Santa Catalina: Fosberg S4516 LAM!; Templeton 11389
LAM!; Trask s.n. in Mar. 1901 NY!; LA; RSA-POM;
SBBG; SBM.

Castilleja foliolosa H. & A.

Santa Catalina: Davidson, A. s.n. Jun. 25, 1891 LAM!;
Beauchamp 349 LAM!; Moran 689 LAM!, NY!; LA;
RSA-POM; SBBG.

Castilleja fruticosa Moran

Guadalupe: Rempel 759-37 LAM!; Moran 12068 LAM!;
Moran 15733 MO!; RSA-POM.

Castilleja grisea Dunkle

San Clemente: Dunkle 7201 LAM!; Trask 183 NY!; El-
more 411 AHFH!; RSA-POM; SBBG.

80 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Castilleja guada/upensis Bdg.

Guadalupe: Palmer 59 MO!, UC!, CM!.

Castilleja hololeuca Greene
San Miguel: RSA-POM; SBBG; SBM.

Santa Rosa: Moran 812 LAM!; RSA-POM; SBBG; SBM.
Santa Cruz: Yates s.n. in Aug. 1893 LAM!; RSA-POM;
SBBG; SBM.

Anacapa: Moran 7 1 5 LAM!; Dunkle 7622 LAM!, AHFH!;
Elmore 226 AHFH!; RSA-POM; SBBG; SBM.
Castilleja mollis Penn.

San Miguel: Schuyler 18 LAM!; Elmore 333 AHFH!; El-
more 341 AHFH!.

Santa Rosa: Munz 11678 POM!, SBM!; Dunkle 8489
AHFH!; Elmore 172 AHFH!; LA; RSA-POM.

Collinsia heterophvlla Buist ex Grah. var. heterophylla
Santa Rosa: SBBG; SBM.

San Clemente: Dunkle 7295 LAM!, AHFH!; Trask 340
NY!; RSA-POM; SBBG.

Diplacus longiflorus Nutt. ssp. longiflorus
Santa Rosa: Sweet s.n. Apr. 15, 1935 POM!.

Santa Cruz: Moran 767 LAM!; Greene s.n. in Jul.-Aug.
1886 ND-G#001718!; Clokey 5206 LA!, NY!, US!; RSA-
POM; SBBG; SBM.

Diplacus parviflorus Greene

Santa Rosa: Dunkle 8439 LAM!, AHFH!; Moran 811
LAM!; Elmore 185 AHFH!; LA; RSA-POM; SBBG;
SBM.

Santa Cruz: Dunkle 8474 LAM!; Fosberg 7533 LAM!; Fos-
berg 7599 LAM!; RSA-POM; SBBG; SBM.

Anacapa: Moran 718 LAM!; Dunkle 7616 LAM!; Dunkle
7658 LAM!; RSA-POM; SBBG; SBM.

San Clemente: RSA-POM; SBBG.

Diplacus puniceus Nutt.

Santa Catalina: Davidson, A. s.n. Jun. 26, 1891 LAM!;
Fosberg S4353 LAM!; Beauchamp 342 LAM!; LA!; RSA-
POM; SBBG.

Galvezia speciosa (Nutt.) Gray
Santa Barbara: Cooper s.n. UC#26766!.

Santa Catalina: Trask s.n. Mar. 1 90 1 LAM!; Fosberg S54 1 5
LAM!; Moran 602 LAM!; RSA-POM; SBBG.

San Clemente: House & Grumbles s.n. USC!; Moran 584
LAM!; Munz 6685 LAM!; RSA-POM; SBBG.
Guadalupe: Rempel 759-37 LAM!; Moran 17423 LAM!,
RSA!; Carlquist 454 RSA!.

Keckiella cordifolia (Benth.) Straw

Santa Rosa: Dunkle 8432 LAM!, AHFH!; Moran 787
LAM!; RSA-POM; SBBG.

Santa Cruz: Dunkle 8554 AHFH!; Ellison s.n. May 12-
15,1929 LA!; Mower s.n. Aug. 7, 1 966 LA!; RSA-POM;
SBBG; SBM.

Anacapa: Dunkle 7656 LAM!, AHFH!.

Santa Catalina: Fosberg S4325 LAM!; Dunkle 1968 LAM!;

Herley s.n. Apr. 26, 1 932 LA!; RSA-POM; SBBG; SBM.
San Clemente: Dunkle 7287 LAM!, AHFH!; RSA-POM;
SBBG.

Linaria bipartita Willd.

Santa Catalina: RSA-POM.

Linaria canadensis (L.) Dum.-Cours. var. lexana (Scheele)
Penn.

San Miguel: RSA-POM.

Santa Rosa: Moran 778 LAM!; RSA-POM; SBBG; SBM.
Santa Cruz: Moran 753 LAM!; Hoffmann s.n. Jun. 15,
1930 LAM!; Fosberg 7652 LAM!, LA!; RSA-POM;
SBBG; SBM.

Anacapa: Moran 731 LAM!; SBM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Moran 695
LAM!; Templeton 11385 LAM!; RSA-POM; SBBG.
San Clemente: Dunkle 7313 LAM!; Dunkle 7210 AHFH!;

RSA-POM; SBBG.

Guadalupe: Moran 17286 RSA!.

Mint ulus brandegei Penn.

Santa Cruz: SBM.

Mimulus brevipes Benth.

Santa Catalina: Dunlavy s.n. May 4, 1934 LA!.

Mimulus cardinalis Dough ex Benth.

Santa Cruz: Dunkle 8646 LAM!; Dunkle 8542 AHFH!;

LA; RSA-POM; SBBG; SBM.

Santa Catalina: Fosberg S4910 LAM!; Fosberg S4846
LAM!; Dunkle 1951 AHFH!; LA; RSA-POM; SBBG.
Mimulus floribundus Dough ex Lindl. var .floribundus
Santa Rosa: SBM.

Santa Catalina: Fosberg 4845 LAM!, NY!; Dunkle 2434
AHFH!; Trask s.n. in Apr. 1896 MO!; RSA-POM.

San Clemente: Trask 341 US!.

Mimulus guttatus Fisch. ex DC. ssp. guttatus

Santa Rosa: Dunkle 8435 LAM!, AHFH!; SBBG; SBM.
Santa Cruz: Fosberg 7645 LAM!; Dunkle 8550 LAM!,
AHFH!; Fosberg 7606 LAM!; RSA-POM; SBBG; SBM.
Santa Catalina: Grant & Wheeler s.n. LAM!; Fosberg 8 1 46
LA!; RSA-POM; SBBG.

San Clemente: Pierson 3741 LA!; RSA-POM; SBBG.
Mimulus guttatus Fisch. ex DC. ssp. littoralis Penn.

Santa Rosa: RSA-POM.

Mimulus guttatus Fisch. ex DC. ssp. micranthus (Heller) Munz
SantaCruz: Hoffmann s.n. Mar. 28, 1925 SBM#788!; Hoff-
mann s.n. May 14, 1927 SBM#8640L
Mimulus latifolius Gray

Guadalupe: Palmer 58 MO!; Greene s.n. Apr. 22, 1885
ND-G!; Palmer 839 US!.

Mimulus nasutus Greene

Santa Cruz: Hoffmann 179 SBM!.

Mimulus traskiae Grant
Santa Catalina: Trask s.n. in Mar. 1901 LAM!.
Orthocarpus attenuatus Gray
Guadalupe: Moran 13821 LAM!, RSA!.

Orthocarpus densiflorus Benth. var. densiflorus
San Miguel: RSA-POM; SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7650 LAM!, LA!; Fosberg 7693 LAM!;

Elmore 459 AHFH!; RSA-POM; SBBG; SBM.

San Nicolas: Trask 61 MO!.

Orthocarpus purpurascens Benth. var. pallidus Keck
San Miguel: RSA-POM.

Santa Rosa: RSA-POM.

Santa Cruz: RSA-POM.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 81

Santa Catalina: Fosberg S4453 LAM!; Fosberg S4938
LAM!; Dunkle 1739 AHFH!.

Orthocarpus purpurascens Benth. var. purpurascens
San Miguel: Schuyler 1 54 LAM!; RSA-POM; SBBG; SBM.
Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Dunkle 2166 AHFH!; Moran 657 LAM!.
Guadalupe: (Keck 1927: Brandegee in 1897 at UC).
Scrophularia californica Cham. & Schlecht. ssp. californica
Santa Rosa: Hoffmann s.n. Apr. 17, 1929 SBM!.
Scrophularia vil/osa Penn, in Millsp. & Nutt.

Santa Catalina: Trask s.n. in Mar. 1901 NY!; Grant 1 186
US!; Fosberg 8133 LA!; RSA-POM; SBBG.

San Clemente: Trask 230 NY!; Raven 17711 LA!; RSA-
POM.

Guadalupe: Moran 8390 RSA!, US!; Moran 5970 RSA!.
Verbascum thapsus L.

Santa Cruz: Dunkle 8555 LAM!, AHFH!; RSA-POM;
SBBG.

Solanaceae
Datura wrightii Regel
Santa Rosa: SBM.

Santa Cruz: Dunkle 8570 LAM!, AHFH!; RSA-POM;
SBBG; SBM.

Santa Catalina: Fosberg S4928 LAM!; RSA-POM; SBBG.
Lycium brevipes Benth. var. brevipes

San Clemente: House & Grumbles s.n. Aug. 5-13, 1930
USC!.

Lycium brevipes Benth. var. hassei (Greene) C.L. Hitchc.
Santa Catalina: Davidson, A. s.n. Jun. 25, 1891 LAM!;
Hasse 4155 NY!; Trask s.n. in Apr. 1896 US!; RSA-
POM.

San Clemente: Trask 331 NY!, US!; Trask 332 GH!, NY!,
US!.

Lycium californicum Nutt.

Anacapa: SBBG.

San Nicolas: Trask s.n. in Apr. 1897 US!; Kanakoff s.n.
Apr. 27, 1940 LAM!; Foreman 123 US!; RSA-POM;
SBBG.

Santa Barbara: Trask s.n. in Apr. 1901 LAM!; Elmore 295
AHFH!; RSA-POM; SBBG.

Santa Catalina: Dunkle 2151 AHFH!; Fosberg S4705
LAM!; Fosberg S4882 LAM!; RSA-POM; SBBG.

San Clemente: Trask 27 US!; Dunkle 7328 LAM!, AHFH!;

Elmore 397 AHFH!; RSA-POM; SBBG.

Guadalupe: Lindsay 43081 RSA!; Moran 5631 RSA!.
Lycium fremontii Gray
Santa Rosa: SBM.

Guadalupe: Moran & Ernst 6733 RSA!.

Lycium verrucosum Eastw.

San Nicolas: Trask 60 LAM!; Trask s.n. in Apr. 1897
CAS#720!.

Nicotiana attenuata Torr. ex Wats, in King

Guadalupe: Palmer 64 NY!; Greene s.n. Apr. 25, 1885
CAS#859!.

Nicotiana bigelovii (Torr.) Wats. var. bigelovii
Santa Catalina: RSA-POM; SBBG.

Nicotiana clevelandii Gray

Santa Cruz: Hoffmann s.n. Jun. 30, 1930 CAS#176935!;

RSA-POM; SBBG; SBM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!.

Nicotiana glauca Grah.

Santa Catalina: Dunkle 1977 AHFH!; Fosberg S4284
LAM!; Fosberg S4577 LAM!; RSA-POM; SBBG.
Guadalupe: Moran 17327 SD!; Wiggins & Ernst 220 DS!;
Moran 2838 DS!.

Petunia parvi flora Juss.

Santa Rosa: Dunkle 8430 LAM!; Elmore 1 70 AHFH!; RSA-
POM; SBBG; SBM.

Solanum douglasii Dunal in DC.

San Miguel: RSA-POM; SBBG; SBM.

Santa Rosa: Dunkle 8450 LAM!; Elmore 200 AHFH!;

Dunn, D. 1304 LA!; RSA-POM; SBBG; SBM.

Santa Cruz: Dunkle 8623 LAM!; AHFH!; Dunkle 8641
AHFH!; Clokey 5201 LA!; RSA-POM; SBBG; SBM.
Santa Catalina: Moran 688 LAM!; Fosberg 4614 LAM!;

Elmore 438 AHFH!; RSA-POM; SBBG.

San Clemente: Dunkle 7225 AHFH!; Elmore 420 AHFH!;

House & Grumbles s.n. USC!; RSA-POM; SBBG.
Guadalupe: Moran 6602 NY!, RSA!; Palmer 61 NY!;
Palmer 6116 RSA!.

Solanum elaeagnifolium Cav.

Santa Cruz: Laughrin 426 SCIR!.

Santa Catalina: Thome 36693 RSA!; SBBG.

Solanum nodiflorum Jacq.

San Nicolas: Dunkle 8324 LAM!, AHFH!; RSA-POM;
SBBG.

San Clemente: RSA-POM.

Guadalupe: Moran 6448 RSA!; Palmer 60 NY!; Palmer
860 NY!.

Solanum sarrachoides Sendt. ex Mart.

Santa Cruz: RSA-POM.

Solanum wallacei (Gray) Parish ssp. clokeyi (Munz) Thome
Santa Rosa: Moran 777 LAM!; Dunn, D. 1305 LA!; Dunn,
N. s.n. May 15, 1932 LA!; SBBG; SBM.

Santa Cruz: Moran 745 LAM!; Elmore 467 AHFH!; Yates
80 LAM!; RSA-POM; SBBG; SBM.

Solanum wallacei (Gray) Parish ssp. wallacei
Santa Catalina: Grant & Wheeler s.n. Apr. 21-26, 1904
LAM!; Fosberg S4285 LAM!; Moran 687 LAM!; LA;
RSA-POM; SBBG.

Guadalupe: Palmer 62 NY!; Franceschi 15 NY!, SBM!;
Moran 18143 SD!.

Tamaricaceae
Tamarix tetrandra Pallas
Santa Cruz: Philbrick B7742 SBBG!.

Anacapa: SBBG.

Tropaeolaceae
Tropaeolum majus L.

Santa Catalina: RSA-POM; SBBG.

Urticaceae

Hesperocnide tenella Torr.

Santa Cruz: Hoffman s.n. SBM#21891!; Hoffman s.n.
SBM#1 1 168!; RSA-POM; SBBG.

82 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Santa Barbara: RSA-POM; SBBG.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!, NY!; Fos-
berg S4347 LAM!; Fosberg S4487 LAM!; RSA-POM.
San Clemente: Trask 188 NY!; RSA-POM; SBBG.
Guadalupe: Palmer 86 NY!.

Parietaria hespera Flinton.

San Miguel: SBBG.

Santa Rosa: Hoffmann s.n. SBM#6276!; RSA-POM.
Santa Cruz: Fosberg 7682 LAM!; Hoffmann s.n.

SBM#1Q26!; Abrams & Wiggins 26 CAS!; RSA-POM.
Anacapa: Hoffmann s.n. Mar. 16, 1929 SBM#5561!; SBBG.
San Nicolas: Trask s.n. in Apr. 1897 US!.

Santa Barbara: Dunkle 7448 LAM!; Dunkle 7422 LAM!,
AHFH!, NY!; Blakley 5624 CAS!; RSA-POM; SBBG.
Santa Catalina: Dunkle 2110 AHFH!; Dunkle 2090 LAM!,
AHFH!; Fosberg S4722 LAM!.

San Clemente: Dunkle 7223 LAM!; Dunkle 7219 LAM!;

Dunkle 7221 AHFH!; RSA-POM; CAS.

Guadalupe: Anthony 240 LAM!, CAS!; Moran 5687 RSA!,
CAS!; Wiggins & Ernst 62 DS!.

Soleirolia soleirolii (Req.) Dandy
San Nicolas: Foreman 120 UC!; RSA-POM; SBBG.
Urtica dioica L. ssp. holoserica (Nutt.) Thome
Santa Cruz: Dunkle 8561 LAM!, AHFH!; Elmore 258
AHFH!; Yates s.n. in Aug. 1893 SBM!; RSA-POM.
Anacapa: Dunkle 7248 LAM!, AHFH!; SBBG.

Santa Catalina: Fosberg S4911 LAM!; Dunkle 1992
AHFH!; RSA-POM.

Urtica urens L.

Santa Rosa: Hoffmann s.n. SBM6277!; RSA-POM; SBBG.
Santa Cruz: Hoffmann s.n. SBM#5416!; Hoffmann s.n.
SBM#1 1856!; RSA-POM.

Santa Catalina: Dunkle 2069 LAM!; Fosberg s.n. Apr. 8,
1931 LAM!; RSA-POM.

Valerianaceae
Centranthus ruber (L.) DC.

Santa Cruz: RSA-POM; SBBG.

Santa Catalina: Fosberg S4418 LAM!; RSA-POM; SBBG.
Verbenaceae

Lippia nodiflora (L.) Michx. var. rosea (D. Don) Munz
Santa Cruz: Daily 519 SCIR!.

Santa Catalina: Fosberg S5414 LAM!.

Verbena bracteata Lag. & Rodr.

Santa Catalina: Fosberg S5403 LAM!; RSA-POM; SBBG.

Verbena lasiostachys Link

Santa Cruz: Dunkle 8643 LAM!, AHFH!; Wolf4160 RSA!;
Clokey 5041 POM!.

Santa Catalina: Fosberg S4843 LAM!, POM!.

San Clemente: Munz 6734 POM!; RSA-POM.

Verbena robusta Greene
Santa Rosa: Thome et al. 48789 RSA!.

Santa Cruz: Dunkle 8548 LAM!, AHFH!; Elmore 291
AHFH!; Thome 36670 RSA!; SBBG; SBM.

Santa Catalina: Fosberg S5412 LAM!; Fosberg S4606
LAM!; Fosberg S4741 LAM!; RSA-POM; SBBG.

Violaceae

Viola pedunculata T. & G. ssp. pedunculata
Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Fosberg 7689 LAM!, LA!; RSA-POM; SBBG;
SBM.

Santa Catalina: Moran 680 LAM!; Fosberg S4455 LAM!;

Dunkle 2080 AHFH!, RSA-POM; SBBG.

San Clemente: Dunkle 7326 LAM!, AHFH!; Moran 572
LAM!; RSA-POM; SBBG.

Viscaceae

Phoradendron bolleanuni ( Seem.) Eichlerssp. densum (Torn.)
Wiens

Guadalupe: Palmer 85 CM!.

Vitaceae

Vitis girdiana Munson

Santa Catalina: Dunkle 1950 LAM!; Fosberg S4827 LAM!;
Knopf & Johnson 1456 DS!; RSA-POM; SBBG.

Monocotyledons

Arecaceae

Erythaea edulis (Wendl.) Wats.

Guadalupe: Franceschi 1 RSA!; Franceschi s.n. 1892 &
1893 US!; Moran 18386 SD!; SBM.

Cyperaceae

Carex barbarae Dewey

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 LAM!; Balls &
Blakley 2362 1 RSA!; Hoffmann 228 POM!; CAS; SBBG;
SBM.

Carex globosa Boott.

Santa Rosa: Thome et al. 48933 RSA!; SBBG; SBM.

Santa Cruz: Hoffmann s.n. Sep. 9, 1931 LAM!; Clokey
4877 LAM!, NY!; Abrams & Wiggins 143 DS!, POM!,
NY; SBBG; SBM; US.

Carex gracilior Mkze.

Santa Rosa: Thome et al. 48709 RSA!; SBM.

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 POM!; SBM.

Carex montereyensis Mkze.

Santa Cruz: Clokey 4874 LAM!; Williams 36 POM!; SBM.

Carex pansa Bailey

Santa Rosa: Blakley 3 1 98 RSA!; Blakley 3200 RSA!, CAS!;
Thorne et al. 49018 RSA!; SBBG; SBM.

Carex praegracilis W. Boott.

Santa Rosa: Munz & Hoffmann 11732 POM!; Hoffmann
718 POM!; Hoffmann s.n. Jun. 13, 1931 RSA!; SBM.

Santa Cruz: Hoffmann s.n. Apr. 13, 1931 RSA!; SBM.

Santa Catalina: Thorne 35902 RSA!, CAS!.

Carex senta Boott.

Santa Cruz: Clokey 4875 LAM!; Williams 44 POM!; Thome
& Everett 36805 RSA!; CAS; SBBG; SBM.

Carex subbracteata Mkze.

Santa Rosa: Hoffmann 717 POM!; Raven, Blakley & Om-
duff 14981 RSA!; SBBG.

Santa Cruz: Hoffmann s.n. Dec. 7, 1930 POM!; Hoffmann
214 POM!; Clokey 4874 POM!.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 83

Carex triquetra Boott.

Santa Cruz: SBM.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!, NY!, US!;
Dunkle 2140 LAM!; Nuttall 131 NY!, US!; CAS; DS;
RSA-POM; SBBG.

Carex tumulicola Mkze.

Santa Rosa: (Smith 1976: Howell 6380 at CAS).

Santa Cruz: Clokey 5971 NY!, UC!.

San Clemente: Raven 18005 CAS!; RSA-POM.

Cyperus alternifolius L.

Santa Cruz: SBM.

Eleocharis macrostachya Britt, in Small

Santa Rosa: Hoffmann s.n. Apr. 8, 1930 SBM!.

San Nicolas: Trask s.n. in 1897 US!; Trask s.n. in Apr.
1901 LAM!; E.Z. Rett & P.C. Orr s.n. in Mar. 1945
SBM#19398!.

Santa Catalina: Trask s.n. in Mar. 1901 LAM!; Trask s.n.

in Mar. 1898 US!; Wolf 3597 US!; RSA-POM; SBBG.
San Clemente: RSA-POM.

Scirpus americanus Pers. var. monophyllus (Presl) Koyama
Santa Rosa: SBM.

Scirpus californicus (C.A. Mey.) Steud.

Santa Cruz: Dunkle 8581 LAM!, AHFH!; Elmore 293
AHFH!; SBBG; SBM.

Scirpus cernuus Vahl. ssp. californicus (Torr.) Thome
San Miguel: RSA-POM; SBM.

Santa Rosa: RSA-POM; SBM.

Scirpus microcarpus Presl

Santa Catalina: RSA-POM; SBBG.

Scirpus olneyi Gray
Santa Rosa: SBM.

San Nicolas: Dunkle 8329 LAM!, AHFH!; Raven &
Thompson 20765 DS!; RSA-POM.

Iridaceae

Chasmanthe aethiopica (L.) N.E. Br.

San Miguel: SBBG.

Iris ochroleuca L.

San Nicolas: SBBG.

Sisyrinchium bellum Wats.

San Miguel: RSA-POM; SBM.

Santa Rosa: Dunkle 8506 AHFH!; RSA-POM; SBM.
Santa Cruz: RSA-POM; SBM.

Santa Catalina: Dunkle 1744 AHFH!; RSA-POM.
Juncaceae

Juncus acutus L. var. sphaerocarpus Engelm.

Santa Catalina: Trask s.n. in Aug. 1901 LAM!; Dunkle
1915 AHFH!; Fosberg S4899 LAM!; RSA-POM.
Juncus balticus Willd.

San Miguel: Dunkle 8408 LAM!, AHFH!; SBBG.

Santa Rosa: SBM.

Santa Cruz: SBM.

Santa Catalina: Fosberg S4619 LAM!; Fosberg S4792
LAM!; RSA-POM; SBM.

Juncus bufonius L.

San Miguel: Dunkle 8406 LAM!; RSA-POM; SBM.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: Fosberg 7636 LAM!; Clokey 4878 LAM!; RSA-
POM; SBM.

San Nicolas: RSA-POM.

Santa Catalina: Fosberg S4751 LAM!; Fosberg S4788
LAM!; Wolf 3608 LAM!; RSA-POM; SBM.

San Clemente: RSA-POM.

Guadalupe: Moran 6663 RSA!; Moran 6646 SD!; Moran
17364 SD!.

Juncus ejfusus L. var. brunneus Engelm.

Santa Cruz: SBBG.

Juncus ejfusus L. var. pacificus Fern. & Wieg.

Santa Cruz: RSA-POM.

Juncus mexicanus Willd.

San Miguel: SBM.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: Daily 702 SCIR!.

Santa Catalina: Thome 34967 RSA!; Thome 37666 RSA!.
Juncus patens E. Mey.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: SBBG; SBM.

San Clemente: RSA-POM.

Juncus phaeocephalus Engelm. var. phaeocephalus
Santa Rosa: RSA-POM.

Juncus textilis Buch.

Santa Catalina: RSA-POM.

Juncus xiphioides E. Mey.

Santa Rosa: SBM.

Santa Cruz: SBM.

Santa Catalina: Fosberg S4861 LAM!; Fosberg S5371
LAM!; Fosberg S4861 LAM!; RSA-POM.

Luzula subsessilis (Wats.) Buch.

Santa Rosa: Moran 806 LAM!; RSA-POM; SBBG; SBM.
Santa Cruz: Wolf 2814 US!; RSA-POM; SBBG; SBM.
Liliaceae

Allium lacunosum Wats. var. lacunosum
Santa Rosa: RSA-POM; SBM.

Santa Cruz: Brandegee s.n. in 1888 NY!.

Allium praecox Bdg.

San Miguel: SBBG.

Santa Rosa: SBM.

Santa Cruz: Hoffmann s.n. Apr. 11, 1931 LAM!; Hoffmann
261 LA!; RSA-POM; SBBG; SBM.

Santa Catalina: Dunkle 1750 AHFH!; Grant & Wheeler
s.n. Apr. 23, 1904 LAM!; Fosberg 689 LAM!; RSA-
POM; SBBG.

San Clemente: Dunkle 7304 AHFH!; Dunkle 7246 AHFH!;
Dunkle 7245 AHFH!; RSA-POM; SBBG.

Asparagus officinalis L.

Santa Catalina: Fosberg S5368 LAM!.

Bloomeria crocea (Torr.) Cov. ssp. crocea
Santa Rosa: Dunn, N. s.n. May 15, 1932 LA#2 1888!; Dunn,
N. s.n. May 15, 1932 LA#21891!; Dunn, N. s.n. May
15, 1932 LA#21902!; RSA-POM; SBBG; SBM.

Santa Cruz: Hoffmann s.n. Apr. 11, 1931 LAM!; Clokey
4879 LAM!; Ellison s.n. May 12-15, 1929 LA!; RSA-
POM; SBBG; SBM.

Santa Catalina: Smith, R.J. s.n. May 24, 1934 LAM!; Fos-
berg S4372 LAM!; Fosberg S4554 LAM!; RSA-POM;
SBBG; USC.

84 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Brodiaea jolonensis Eastw.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: SBBG.

Santa Catalina: RSA-POM.

Brodiaea kinkiensis Niehaus
San Clemente: RSA-POM.

Calochortus albus Dougl. ex Benth. var. a/bus
Santa Rosa: Hoffmann s.n. Jun. 6, 1930 LAM!; Dunn, N.

s.n. May 15, 1932 LA!; RSA-POM; SBBG.

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 LAM!; Clokey
4882 LAM!, NY!; Ellison s.n. May 12-15, 1929 LA!;
RSA-POM; SBBG.

Santa Catalina: Brown, C. s.n. May 31, 1925 USC!.
Calochortus catalinae Wats.

Santa Rosa: Moran 781 LAM!, NY!; RSA-POM; SBBG;
SBM.

Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Fosberg S4649 LAM!; Dunkle 1762
AHFH!; Fosberg 8167 NY!; LA; RSA-POM; SBBG.
Calochortus luteus Dougl. ex Lindl.

Santa Cruz: RSA-POM; SBBG; SBM.

Calochortus splendens Dougl. ex Benth.

Santa Catalina: Fosberg S4690 LAM!; Hasse 2753 LAM!;
Trask s.n. in Mar. 1896 NY!; RSA-POM.

Chlorogalum pomeridianum (DC.) Kunth
Santa Rosa: RSA-POM; SBM.

Santa Catalina: RSA-POM.

Dichleostemma pulchellum (Salisb.) Heller
San Miguel: RSA-POM; SBM.

Santa Rosa: Dunn, N. s.n. May 15, 1932 LA!; RSA-POM;
SBM.

Santa Cruz: Clokey 488 1 LAM!; Fosberg 7633 LAM!, LA!;

RSA-POM; SBM.

Anacapa: SBBG; SBM.

San Nicolas: Trask 87 LAM!; Kanakoff s.n. Apr. 12, 1940
LAM!; RSA-POM; SBM.

Santa Barbara: Dunkle 7419 LAM!, AHFH!; RSA-POM;
SBM.

Santa Catalina: Fosberg S4632 LAM!; Fosberg S4369
LAM!; Templeton 11398 LAM!; RSA-POM; USC.

San Clemente: Moran 580 LAM!; Dunkle 7304 LAM!;

Elmore 391 AHFH!; LA; RSA-POM.

Guadalupe: Moran 6651 SD!; Moran 17415 SD!; Moran
20306 SD!.

Lilium humboldtii Roezl & Leichtl. ssp. ocellatum (Kell.)
Thome

Santa Rosa: RSA-POM.

Santa Cruz: Clokey 4885 LAM!; Clokey 4884 LAM!; Hoff-
mann s.n. Apr. 1 1, 1930 LAM!; LA; RSA-POM; SBBG;
SBM.

Triteleia Clementina Hoov.

San Clemente: Dunkle 7283 LAM!, AHFH!; Dunkle 7324
LAM!; Moran 702 LAM!; RSA-POM; SBBG.

Triteleia guada/upensis Lenz
Guadalupe: Moran 12063 RSA!, SD!.

Zigadenus fremontii Torr. var. fremontii
Santa Rosa: Munz & Crow 11616 LA!; RSA-POM; SBBG;
SBM.

Santa Cruz: Fosberg 7529 LAM!; RSA-POM; SBBG; SBM.
Anacapa: Moran 716 LAM!; SBBG; SBM.

Orchidaceae

Epipactis gigantea Dougl. ex Hook.

Santa Cruz: RSA-POM; SBM.

Habenaria elegans (Lindl.) Boland.

Santa Rosa: Hoffmann s.n. Jun. 13, 1930 CAS#167564!;

Rowntree s.n. Jun. 16, 1970 CAS#297877!; SBBG; SBM.
Santa Cruz: Clokey 4886 LAM!; Clokey 4887 LAM!; Hoff-
mann s.n. Jun. 14, 1930 CAS#176936!; SBBG; SBM.
Santa Catalina: SBBG.

Habenaria unalascensis (Spreng.) Wats.

Santa Cruz: RSA-POM.

Santa Catalina: Fosberg S4495 LAM!; Fosberg S4568
LAM!; Fosberg S4867 LAM!.

Poaceae

Agrostis diegoensis Vasey

Santa Rosa: Brandegee, T.S. 66 US!; RSA-POM; SBBG;
SBM.

Santa Cruz: Hoffmann s.n. May 23, 1932 POM# 180275!;
SBBG; SBM.

Santa Catalina: Fosberg S4729 LAM!; Fosberg S4869
LAM!; Brandegee, T.S. 50 US!; RSA-POM; SBBG.

San Clemente: Abrams & Wiggins 370 DS!, US!.

Agrostis exarata Trin.

Santa Rosa: Hoffmann s.n. Aug. 7, 1930 SBM!; SBM.
Santa Cruz: RSA-POM; SBBG; SBM.

Santa Catalina: Thome & Thorne 36434 RSA!.

Agrostis semiverticillata (Forsk.) C. Chr.

Santa Rosa: Thome et al. 48960 RSA!; SBBG; SBM.
Santa Cruz: Dunkle 8553 LAM!, AHFH!, RSA!; Hoffmann
s.n. Jun. 28, 1930 LAM!; Dunkle 8635 LAM!, RSA!;
SBBG; SBM.

Anacapa: Dunkle 7650 LAM!.

Santa Catalina: Trask s.n. in Mar. 1901 NY!; Thome 36636
RSA!; Raven 17844 RSA!; SBBG.

Ammophila arenaria (L.) Link

San Nicolas: Foreman 101 UC!; Wier & Beauchamp s.n.
Jul. 2, 1978 RSA!.

Andropogon glomeratus (Walt.) BSP.

Santa Cruz: Benedict s.n. Nov. 16, 1969 SBBG.

Aristida adscensionis L. var. modesta Hack, in Stuckert
Santa Rosa: SBBG.

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 LAM!; Thorne
& Everett 36806 RSA!; Raven & Smith 15201 RSA!,
DS!; SBM.

Santa Catalina: Brandegee 56 US!; Thome & Thome 42473
RSA!; Thome 35942 RSA!.

San Clemente: Abrams & Wiggins 364 DS!, GH!; Raven
17609 RSA!; Raven 17650 RSA!.

Guadalupe: Wiggins & Ernst 215 SD!; Palmer 675 US!.

Aristida divaricata Humb. & Bonpl. in Willd.

Santa Cruz: SBBG.

Arundo donax L.

San Nicolas: Beauchamp s.n. Jul. 1, 1978 RSA!; SBBG.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 85

Avena barbata Brot.

San Miguel: SBBG.

Santa Rosa: Dunkle 8523 LAM!, AHFH!; Dunn, N. s.n.
May 15, 1932 LA# 11004!; RSA-POM; SBBG; SBM.

Santa Cruz: Dunkle 8565 LAM!, AHFH!; Clokey 4861
LAM!; Fausett 16 LA!; RSA-POM; SBBG; SBM.

Anacapa: SBBG; SBM.

San Nicolas: Kanakoff s.n. LAM!; Foreman & Smith 166
LA!; Thome et al. 52362 RSA!.

Santa Barbara: RSA-POM.

Santa Catalina: Fosberg S4298 LAM!; Detmers s.n. Apr.
14, 1929 USC!; Thome 35893 RSA!.

San Clemente: Dunkle 7255 LAM!, AHFH!; Thome 36 1 45
RSA!; Raven 17290 RSA!.

Guadalupe; Moran 5616 SD!; Palmer 668 ND-G!; Mason
1542 CAS!.

Avena fatua L.

San Miguel: Dunkle 8363 LAM!, AHFH!; SBBG.

Santa Rosa: Thome et al. 48953 RSA!; SBM.

Santa Cruz: SBBG; SBM.

Anacapa: SBBG.

San Nicolas: Raven & Thompson 20776 DS!; Trask 17
US!.

Santa Barbara: Dunkle 7454 LAM!, AHFH!; Philbrick
B6846 RSA!; SBM.

Santa Catalina: Dunkle 1792 AHFH!; Trask s.n. in May
1898 US!; Thome 33452 RSA!.

San Clemente: Trask 242 US!; Munz 6619 POM!; Raven
17133 RSA!.

Guadalupe: Palmer 94 CM!; Brandegee, T.S. s.n. May 24,
1897 UC!.

Avena sativa L.

Santa Rosa: SBM.

Santa Cruz: SBBG.

Santa Catalina: Fosberg S4425 LAM!; Fosberg S4662
LAM!; Thome 36274 RSA!.

San Clemente: Raven 17622 RSA!.

Bothriochloa barbinodis (Lag.) Herter

Santa Catalina: McClatchie s.n. Sep. 1 2, 1 893 NY!; Thome
36196 RSA!; Wolf 4026 RSA!.

Brachypodium distachyon (L.) Beauv.

Santa Catalina: Fosberg S4964 LAM!, RSA!, SBM!; Thome
36891 RSA!; Raven 17825 LA!, RSA!.

Bromus arizonicus (Shear) Steb.

San Miguel: Hoffmann s.n. Apr. 10, 1930 US!; Beck, R.H.
1 US!.

Santa Rosa: Hoffmann 706 SBM!; Hoffmann 727 SBM!;
Hoffmann s.n. Apr. 20, 1929 SBM#7652!, CAS# 168467!.

Santa Cruz: (Stebbins, Tobgy & Harlan 1944: Brandegee
s.n. in 1888 at UC).

Anacapa: Dunkle 7445 LAM!; SBBG.

San Nicolas: Howell 8226 CAS!, US!.

Santa Barbara: Dunkle 7455 LAM!, AHFH!; Philbrick &
Ricker B69-55 US!; Piehl 63132 CAS!; RSA-POM.

Santa Catalina: Fosberg S4646 LAM!; Millspaugh 4632
US!; Brandegee, T.S. 57 US!; RSA-POM.

San Clemente: Trask 357 NY!, US!; RSA-POM.

Bromus carinatus H. & A.

San Miguel: Beck, R.H. 2 US!; Hoffmann 728 SBM!.
Santa Rosa: Hoffmann s.n. Apr. 18, 1929 SBM#7649!;
Hoffmann s.n. Jun. 12, 1930 SBM#10181!; Hoffmann
s.n. Apr. 8, 1930 US# 1535533!; RSA-POM.

Santa Cruz: Hoffmann 262 LAM!; Hoffmann s.n. Apr. 12,
1931 LAM!, SBM!; Hoffmann s.n. Mar. 22, 1929
SBM#5454!; RSA-POM.

Anacapa: SBBG; SBM.

San Nicolas: Trask 13 US!; Trask 14 US!; RSA-POM.
Santa Catalina: Fosberg S4608 LAM!; Fosberg S4484
LAM!; Trask s.n. in May 1897 US!; RSA-POM.

San Clemente: RSA-POM.

Bromus diandrus Roth.

San Miguel: Dunkle 8361 LAM!; Bond 405 SBM!; Bond
406 SBM!; SBBG.

Santa Rosa: Hoffmann s.n. Apr. 20, 1929 SBM#7595!;

Hoffmann s.n. Apr. 20, 1929 SBM#7648!; RSA-POM.
Santa Cruz: Hoffmann s.n. Apr. 12, 1931 SBM#11160!;

Clokey 4856 LAM!; Dunkle 8564 LAM!; LA; RSA-POM.
Anacapa: Bond 329 SBM!; Bond 339 SBM!; SBBG.

San Nicolas: Kanakoff s.n. Apr. 16, 1940 LAM!; Blakley
4078 US!; Foreman & Smith 155 LA!; RSA-POM.
Santa Barbara: Dunkle 8 1 39 LAM!; AHFH!; Dunkle 8141
LAM!; AHFH!; RSA-POM.

Santa Catalina: Dunkle 1787 LAM ! ; Fosberg S4 3 5 6 LAM ! ;

Detmers s.n. Apr. 14, 1929 LAM!; RSA-POM.

San Clemente: Piehl 62397 US!; RSA-POM.

Guadalupe: Moran 13773 LAM!, RSA!, SD!; Wiggins &
Ernst 77 DS!, SD!; Brown, W.W. 31 GH!.

Bromus madritensis L.

Santa Cruz: Fausett 23 LA!; RSA-POM.

Bromus maritimus (Piper) Hitchc.

San Miguel: Hoffmann 681 SBM!; Bond, R. s.n. Apr. 18,
1939 SBM!; Hoffmann s.n. Apr. 10, 1930 SBM#9294!;
RSA-POM.

Santa Rosa: Hoffmann 724 SBM!.

Santa Cruz: Yates s.n. in Aug. 1893 SBM# 15947!; SBBG.
Anacapa: Dunkle 7634 LAM!; [Hoffmann] s.n. Mar. 11,
1928 SBM#4136!; SBBG.

San Nicolas: Philbrick & Benedict B69-161 US!.

Santa Catalina: Fosberg S4484 SBM!.

Bromus mollis L.

San Miguel: SBBG.

Santa Rosa: Hoffmann 744 SBM!; Dunkle 8520 LAM!;

Hoffmann s.n. Apr. 15, 1929 SBM#6278!; RSA-POM.
Santa Cruz: Hoffmann s.n. Jun. 15, 1930 SBM#2304!;

Clokey 4855 LAM!, US!; Fausett 17 LA!; RSA-POM.
Anacapa: Hoffmann s.n. Mar. 1 1, 1928 SBM#4099!; Bond
332 SBM!; SBBG.

San Nicolas: Kanakoff s.n. Apr. 19, 1940 LAM!; Dunkle
8308 LAM!, AHFH!; P.C. Orr & E.Z. Rett s.n. Mar. 26,
1945 SBM#19343!; RSA-POM.

Santa Barbara: RSA-POM.

Santa Catalina: Fosberg S4493 LAM!; Fosberg S4566
LAM!; Dunkle 1916 AHFH!; RSA-POM; USC.

San Clemente: Dunkle 7254 LAM!, AHFH!; Dunkle 7362

86 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

AHFH!; E.Z. Rett & P.C. Orr s.n. Apr. 10, 1945
SBM#19338!; RSA-POM.

Guadalupe: Wiggins & Ernst 105 DS!, SD!; Moran 6642
SD!; Moran 2881 DS!.

Bromus pseudolaevipes Wagnon.

Santa Rosa: [no collector, probably Hoffmann] s.n. Jun.
13, 1930 SBM#10180!; [no collector, probably Hoff-
mann] s.n. Jun. 12, 1930 SBM#10182!; RSA-POM.
Santa Cruz: Hoffmann s.n. Jun. 29, 1930 SBM#10132!,
US# 1649447!; SBBG.

Santa Catalina: Fosberg S450 1 LAM!, SBM!; Trask s.n. in
Mar. 1901 NY!; Raven 17772 US!; RSA-POM; SBBG.
Bromus rubens L.

San Miguel: SBBG.

Santa Rosa: Dunkle 8519 LAM!; Hoffmann s.n. May 6,
1932 SBM!; RSA-POM; SBBG.

Santa Cruz: Hoffmann s.n. Apr. 12, 1931 LAM!, SBM!;
Dunkle 8639 LAM!, AHFH!; Fausett 1 8 LA!; RSA-POM;
SBBG.

Anacapa: Bond 330 SBM!; SBBG.

San Nicolas: RSA-POM.

Santa Barbara: Dunkle 7423 LAM!, AHFH!; Bond 373
SBM!; Bond 395 SBM!; RSA-POM.

Santa Catalina: Fosberg S4296 LAM!; Dunkle 1778
AHFH!; Dunkle 1917 AHFH!; RSA-POM.

San Clemente: Dunkle 7256 LAM!; RSA-POM.
Guadalupe: Moran 5976 SD!; Wiggins & Ernst 76 DS!;
Moran 12026 RSA!, SD!.

Bromus stamineus Desv. in Gray
Santa Cruz: Raven & Smith 15290 RSA!.

Bromus sterilis L.

Santa Catalina: Nuttall 95 F!, US!; Nuttall 562 F!; Nuttall
1219 F!.

Bromus tectorum L.

Guadalupe: Palmer 99 (in part) NY!.

Bromus trinii Desv. in C. Gay

Santa Rosa: [no collector, probably Hoffmann] s.n. Apr.
8, 1930 SBM#10184!.

Santa Cruz: Hoffmann s.n. Apr. 22, 1932 SBM# 11947!;

Brandegee s.n. in 1888 (in part) UC#121663!.

Anacapa: [no collector, probably Hoffmann] s.n. Mar. 1 1,
1928 SBM#409 1 ! (in part).

San Nicolas: Trask 1 MO!.

Santa Barbara: Dunkle 7424 LAM!; Piehl 63151 RSA!.
Santa Catalina: Brandegee s.n. Mar. 12, 1890 UC!; Trask
s.n. in May 1897 MO!.

San Clemente: Raven 17185 RSA!.

Guadalupe: Palmer 667 NY!, UC!, US!; Palmer 658 US!;
Moran 17369 SD!; CSA; DS.

Calamagrostis rubescens Buckl.

Santa Cruz: Blakley & Muller 3726 SCIR!; SBBG.
Cortaderia atacamensis (Phil.) Pilger
Santa Cruz: SBM.

Santa Catalina: RSA-POM.

Crypsis aculeata (L.) Ait.

Santa Catalina: RSA-POM.

Cynodon daclylon (L.) Pers.

Santa Rosa: Dunkle 8474 LAM!; RSA-POM; SBM.

Santa Cruz: Dunkle 8652 LAM!; RSA-POM; SBM.
Anacapa: SBBG.

San Nicolas: Dunkle 8325 LAM!; Trask s.n. in Apr. 1897
NY!; RSA-POM.

Santa Catalina: Fosberg S4439 LAM!; Nuttall 540 US!;
RSA-POM.

San Clemente: RSA-POM.

Dactylis glomerata L.

Santa Catalina: RSA-POM.

San Clemente: SBBG.

Deschampsia danthomoides (Trin.) Munro in Benth.

San Clemente: RSA-POM.

Dissanthelium californicum (Nutt.) Benth.

Santa Catalina: Gambel s.n. GH!; MO!.

San Clemente: Trask 324 US!.

Guadalupe: Palmer 96 CM!, MO!, NY!.

Distichlis spicata (L.) Greene var. stolonifera Beetle
San Miguel: Dunkle 8366 LAM!; Hoffmann s.n. Apr. 10,
1930 SBM#9367!; Bond 416 SBM!; SBBG.

Santa Rosa: Dunkle 8526 LAM!, AHFH!; Hoffmann s.n.

Apr. 14, 1929 SBM!; RSA-POM; SBBG.

Santa Cruz: Dunkle 8587 LAM!, AHFH!; Clokey 4859
LAM!; Elmore 289 AHFH!; RSA-POM; SBBG; SBM.
Anacapa: Bond 353 SBM!; SBBG.

San Nicolas: Dunkle 8314 LAM!, AHFH!; Kanakoff s.n.
Apr. 19, 1940 LAM!; Trask s.n. in Apr. 1901 LAM!,
US!; LA; RSA-POM; SBM.

Santa Catalina: Trask s.n. in Sep. 1 896 US!; Fosberg S4475
LAM!; Fosberg S4871 LAM!, SBM; RSA-POM.

San Clemente: Trask s.n. Oct. 1902 US!; Dunkle 7331
LAM!, AHFH!.

Echinochloa crus-galli (L.) Beauv. var. crus-galli
Santa Catalina: Thorne 36427 RSA!.

Ehrharta calycina Sm.

Santa Catalina: Thorne, Rollins, Propst & Carolin 36741
RSA!.

San Clemente: SBBG.

Elymus condensalus Presl

San Miguel: Dunkle 8371 LAM!, AHFH!; RSA-POM;
SBM.

Santa Rosa: SBM.

Santa Cruz: Clokey 4867 LAM!; RSA-POM; SBM.
Anacapa: SBBG; SBM.

Santa Catalina: Johnstone s.n. May 20, 1934 USC!; RSA-
POM.

San Clemente: Murbarger 59 UC!.

Elymus glaucus Buckl. ssp. glaucus

Santa Cruz: Hoffmann 598 US!; RSA-POM; SBM.

Santa Catalina: Fosberg S4610 LAM!; Fosberg S4853
LAM!, NY!; Brandegee 49 US!; RSA-POM.

Elymus pacificus Gould
San Miguel: Bond 419 SBM!.

Elymus triticoides Buckl.

San Miguel: Dunkle 8362 LAM!; Elmore 3 1 7 AHFH!; RSA-
POM; SBBG.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 87

Santa Rosa: Dunkle 8518 LAM!, AHFH!; Dunn, N. s.n.
May 15, 1932 LA!; SBM.

Santa Cruz: Ellison, L. s.n. LA# 14864!; RSA-POM; SBBG;
SBM.

Anacapa: SBBG.

Santa Catalina: Fosberg S4831 LAM!; Trask s.n. in Mar.
1901 NY!; Trask s.n. in May 1898 US!; RSA-POM.
Festuca arundinacea Schreb.

Santa Catalina: RSA-POM.

Gastridium ventricosum (Gouan) Schinz & Thell.

Santa Rosa; SBM.

Santa Cruz: Clokey 4868 LAM!; Clokey 4869 LAM!; Hoff-
mann s.n. Apr. 12, 1931 LAM!, LA!; RSA-POM; SBM.
Santa Catalina: RSA-POM.

San Clemente: Dunkle 7348 LAM!, AHFH!; RSA-POM.
Hordeum californicum Covas & Steb.

San Miguel: Blakley 5837 DS; SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: RSA-POM.

Anacapa: SBBG; SBM.

San Nicolas: Trask s.n. in Apr. 1901 LAM!; RSA-POM;
SBBG.

Santa Catalina: Fosberg S4945 LAM!; RSA-POM.
Hordeum depressum (Scribn. & Sm.) Rydb.

Santa Cruz: Abrams & Wiggins 125 DS!.

Hordeum geniculatum All.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: Clokey 5166 LA!, NY!, RSA!.

Santa Catalina: RSA-POM.

San Clemente: RSA-POM.

Hordeum murinum L. ssp. glaucum (Steud.) Tzvel.

San Miguel: Dunkle 8365 LAM!, AHFH!; SBM.

Santa Rosa: SBBG; SBM.

Santa Cruz: Clokey 4858 LAM!, NY!; Raven & Smith
15146 CAS!; SBBG; SBM.

Anacapa: SBBG; SBM.

San Nicolas: Trask s.n. in Apr. 1901 NY!; Howell 8211
CAS!; Raven & Thompson 20725 DS!; RSA-POM.
Santa Barbara: Bryan, Dr. & Mrs. s.n. Jul. 14, 1922 LAM!;
Dunkle 8108 LAM!, AHFH!, DS!, NY!; Blakley 5673
DS!; RSA-POM.

Santa Catalina: Dunkle 1906 AHFH!; Knopf 28 F!; Trask
s.n. in Mar. 1901 NY!; RSA-POM.

San Clemente: Trask 259 NY!; Meams 4055 DS!; RSA-
POM.

Guadalupe: Palmer 671 F!; Webber & McCoy 1 1960 DS!;
Moran 17386 SD!.

Hordeum murinum L. ssp. leporinum (Link) Arcangeli
San Miguel: Dunkle 8365 LAM!, AHFH!; Munz & Crow
11815 LA!; Hoffmann s.n. Apr. 10, 1930 CAS#177736!;
SBBG.

Santa Rosa: Dunn, N. s.n. May 15, 1932 LA!; RSA-POM;
SBBG.

Santa Cruz: Clokey 4857 LAM!; Fausett 14 LA!; SBBG;
SBM.

Anacapa: Blakley 2748 CAS!; SBBG; SBM.

San Nicolas: Kanakoff s.n. Apr. 19, 1940 LAM!; Foreman
& Lloyd 132 LA!; SBBG.

Santa Catalina: Fosberg S4295 LAM!; Dunkle 1784
AHFH!; Millspaugh 4660 F!; DS; RSA-POM.

San Clemente: House & Grumbles s.n. Aug. 5-13, 1930
USC!; Dunkle 7271 LAM!; AHFH!; RSA-POM.
Guadalupe: Wiggins & Ernst 12, DS!, SD!; Moran 17321
SD!; Mason 1544 F!.

Hordeum pusillum Nutt.

San Miguel: SBBG.

Santa Cruz: RSA-POM.

Anacapa: SBBG; SBM.

Santa Barbara: RSA-POM.

Santa Catalina: Fosberg S4666 LAM!; Fosberg S4706
LAM!; RSA-POM; SBM.

San Clemente: RSA-POM.

Hordeum vulgare L.

Santa Rosa: SBBG.

Santa Cruz: Raven & Smith 15287 CAS!; SBBG.

Santa Catalina: Fosberg S4567 LAM!; Fosberg S4512
LAM!; RSA-POM.

San Clemente: RSA-POM.

Koeleria pyramidata (Lam.) Beauv.

Santa Rosa: Dunkle 8495 LAM!, AHFH!; RSA-POM; SBM.
Santa Cruz: RSA-POM.

Lamarckia aurea (L.) Moench
San Miguel: SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Dunkle 8549 LAM!; RSA-POM; SBBG; SBM.
Anacapa: SBBG.

San Nicolas: Kanakoff s.n. LAM!; RSA-POM.

Santa Barbara: Dunkle 8124 LAM!, AHFH!; Dunkle 7458
LAM!, AHFH!; RSA-POM.

Santa Catalina: Fosberg S4297 LAM!; Fosberg S4660
LAM!; Dunkle 1702 AHFH!; RSA-POM.

San Clemente: Dunkle 7332 LAM!, AHFH!; RSA-POM.
Lolium perenne L. ssp. multiflorum (Lam.) Husnot
Santa Cruz: RSA-POM; SBM.

San Nicolas: Foreman, Evans & Rainey 75 LA!; SBBG;
SBM.

Santa Catalina: Fosberg S4966 LAM!; RSA-POM; SBBG.
San Clemente: RSA-POM; SBBG.

Lolium perenne L. ssp. perenne

Santa Cruz: Dunkle 8634 LAM!; SBBG.

San Nicolas: Foreman 1 1 1 UC!.

Santa Catalina: SBBG.

San Clemente: RSA-POM.

Lolium strictum Presl

Santa Catalina: RSA-POM.

Lolium temu/entum L.

Santa Rosa: SBM.

Santa Cruz: Hoffmann s.n. Apr. 11, 1931 LAM!; Hoffmann
s.n. Apr. 12, 1931 LAM!; SBM.

Santa Catalina: Fosberg S4482 LAM!; Fosberg S4661
LAM!; Brandegee s.n. May 26, 1890 US!; RSA-POM.
San Clemente: Trask 241 US!; RSA-POM.

Melica imperfecta Trin.

San Miguel: Blakley 5839 DS!; RSA-POM; SBBG; SBM.
Santa Rosa: Thome et al. 48945 RSA!; SBBG; SBM.
Santa Cruz: Fosberg 7609 LAM!, LA!; Clokey 4872 LAM!,

88 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

US!; Abrams & Wiggins 15 CAS!, DS!; RSA-POM;
SBBG; SBM.

Anacapa: SBBG; SBM.

Santa Barbara: Dunkle 7438 LAM!, AHFH!; Piehl 63 1 52a
US!; Blakley 5611 DS!; RSA-POM; SBBG.

Santa Catalina: Fosberg S5463 LAM!; Nuttall 350 NY!;

Chase 5567 US!; RSA-POM; SBBG; SBM; USC.

San Clemente: House & Grumbles s.n. USC!; Trask 325
US!; Trask 358 US!; RSA-POM; SBBG.

Guadalupe: Moran 13783 SD!, DS!, US!; Wiggins & Ernst
102 DS!, SD!; Moran 13791 RSA!, SD!.

Monanthochloe littoralis Engelm.

San Miguel: Elmore 318 AHFH!.

Santa Rosa: Dunkle 8491 AHFH!; RSA-POM; SBBG.
Santa Cruz: Hoffmann s.n. Mar. 24, 1929 (in part)
SBM#55 10!.

Santa Catalina: Fosberg S4927 LAM!; Blakley 4729 US!;
Pendleton & Reed 1425 US!.

Muhlenbergia microsperma (DC.) Kunth
Santa Rosa: SBM.

Santa Cruz: Fosberg 7635 LAM!; RSA-POM; SBBG; SBM.
Anacapa: SBM.

Santa Barbara: Dunkle 7421 LAM!, AHFH!; Piehl 63155
US!; RSA-POM; SBBG.

Santa Catalina: Trask G12 in Oct. 1896 US!; Trask s.n. in
Mar. 1901 US!; Nuttall 323 US!; RSA-POM; SBBG.
San Clemente: Blakley 6325 US!; RSA-POM; SBBG.
Guadalupe: Palmer 656 ND-G!; Palmer 670 ND-G!; Mo-
ran 17338 SD!.

Oryzopsis miliacea (L.) Benth. & Hook, ex Aschers. &
Schweinf.

Santa Rosa: SBM.

Santa Cruz: Daily 432 SCIR!.

Santa Catalina: RSA-POM; SBBG.

Parapholis incurva (L. f.) C.E. Hubb.

San Miguel: SBBG.

Santa Rosa: Hoffmann s.n. Apr. 9, 1930 US!; RSA-POM;
SBBG; SBM.

Santa Cruz: RSA-POM; SBBG.

San Nicolas: Kanakoff s.n. Apr. 14, 1940 LAM!; Foreman
179 US!; Blakley 4008 US!; RSA-POM.

Santa Barbara: Philbrick B69-89 SBBG; Philbrick B69-82
SBBG.

Santa Catalina: Fosberg S4946 LAM!; Dunkle 1908
AHFH!; Brandegee 54 US!; DS; RSA-POM.

San Clemente: Raven 17276 US!; RSA-POM.

Paspalum dilitatum Poir. in Lam.

Santa Catalina: RSA-POM.

Paspalum distichum L.

Santa Catalina: RSA-POM.

Pennisetum clandestinum Hochst. ex Chiov.

Santa Cruz: Cox, W. s.n. Nov. 20, 1975 SCIR#0384!.

Phalaris aquatica L.

Santa Catalina: Blakley 5423 US!; RSA-POM.

Phalaris canariensis L.

Santa Catalina: Fosberg S4474 LAM!.

Phalaris caroliniana Walt.

Santa Cruz: Brandegee s.n. in Jun. 1888 UC!.

San Nicolas: Trask 9 MO!.

Santa Barbara: Bryan, Dr. & Mrs. s.n. Jul. 14, 1922 LAM!.
Santa Catalina: Trask s.n. in Mar. 1901 MO!.

San Clemente: RSA-POM.

Guadalupe: Moran 6683 RSA!, SD!; Wiggins & Ernst 125
DS!; Moran 17368 SD!.

Phalaris lemmonii Vasey
Santa Rosa: SBM.

Santa Catalina: Trask s.n. May 1897 MO!; Trask s.n. Mar.
1901 NY!.

San Clemente: Thorne 42957 MO!.

Phalaris minor Retz.

San Miguel: Piehl 6253 DS!; SBBG; SBM.

Santa Rosa: SBM.

Santa Cruz: Hoffmann s.n. Apr. 13, 1931 LAM!; Hoffman
s.n. Jun. 15, 1930 SBM#7556!; Raven & Smith 15142
CAS!; RSA-POM.

Anacapa: SBM.

San Nicolas: Raven & Thompson 20792 DS!; RSA-POM.
Santa Barbara: Blakley 5680 CAS!; RSA-POM.

Santa Catalina: Fosberg S4595 LAM!; Wolf 358 1 DS!; RSA-
POM.

San Clemente: Thome 42891 MO!; RSA-POM.
Guadalupe: Moran 6789 SD!; Wiggins & Ernst 216 DS!.
Phalaris paradoxa L.

San Clemente: RSA-POM.

Poa annua L.

San Miguel: Hoffmann s.n. Apr. 19, 1932 SBM!.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: RSA-POM; SBM.

Santa Catalina: RSA-POM.

San Clemente: RSA-POM.

Guadalupe: Moran 6622 RSA!, SD!; Howell 8260 CAS!;
Moran 25380 SD!.

Poa bolanderi Vasey ssp. howellii (Vasey & Scribn.) Keck
Santa Cruz: Hoffmann s.n. Apr. 12, 1931 SBM#1 1 158!.
Poa douglasii Nees
San Miguel: RSA-POM; SBM.

Santa Rosa: RA-POM; SBM.

Poa palustris L.

Santa Catalina: Trask G-4 in Mar. 1897 MO!.

Poa scabrella (Thurb.) Benth. ex Vasey
Santa Rosa: RSA-POM; SBM.

Santa Cruz: Fosberg 7653 LAM!; Fosberg 7608 LAM!;

RSA-POM; SBM.

Anacapa: SBBG; SBM.

Santa Catalina: Fosberg S4521 LAM!; Trask s.n. in Mar.

1901 LAM!, MO!; Fosberg S4653 LAM!; RSA-POM.
San Clemente: RSA-POM.

Guadalupe: Moran 13816 RSA!, SD!.

Polypogon interrupt us HBK.

San Miguel: RSA-POM; SBM.

Santa Rosa: Dunkle 8502 LAM!; SBBG; SBM.

Santa Cruz: Clokey 4860 LAM!; Hoffmann 48 LAM!; Dun-
kle 8551 LAM!; RSA-POM; SBBG; SBM.

Santa Catalina: Fosberg S4511 LAM!; Dunkle 1980

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 89

AHFH!; Brandegee s.n. May 13, 1890 UC!; RSA-POM;
SBBG.

San Clemente: Murbarger 239 UC!.

Polypogon monspeliensis (L.) Desf.

San Miguel: Dunkle 8395 LAM!; RSA-POM; SBM.

Santa Rosa: Elmore 193 AHFH!; Hoffmann s.n. Apr. 9,
1930 CAS#19 1 780!; RSA-POM; SBM.

Santa Cruz: Dunkle 8552 LAM!; Hoffmann s.n. Jun. 28,
1930 (in part) LAM!; Elmore 267 AHFH!; RSA-POM;
SBM.

San Nicolas: Trask s.n. in Mar. 1901 LAM!; Dunkle 8327
LAM!, AHFH!; E.Z. Rett & P.C. Orr s.n. SBM!; DS;
LA; RSA-POM.

Santa Barbara: Dunkle 8148 AHFH!; Bryan, Dr. & Mrs.

s.n. Jul. 14, 1922 LAM!; RSA-POM.

Santa Catalina: Trask s.n. in May 1901 LAM!; Fosberg
S4473 LAM!; Dunkle 1912 AHFH!; RSA-POM.

San Clemente: RSA-POM.

Guadalupe: Rempel 758-37 LAM!; Moran 6604 RSA!,
SD!; Moran 17301 SD!.

Schismus arabicus Nees
Santa Catalina: RSA-POM.

Schismus barbatus Thell.

Santa Cruz: Daily 607 SCIR#0814!.

Scleropoa rigida (L.) Griseb.

Santa Catalina: RSA-POM.

Sitanion jubatum J.G. Sm.

Santa Catalina: Nuttall 314 F!.

Sorghum bicolor (L.) Moench
Santa Catalina: RSA-POM.

Sorghum halepense (L.) Pers.

Santa Catalina: Fosberg S4513 LAM!; Dunkle 2464
AHFH!.

Stipa cernua Steb. & Love
Santa Rosa: Hoffmann s.n. CAS!; RSA-POM.

Santa Cruz: Breedlove 2879 DS!; Abrams & Wiggins 216
DS!; RSA-POM.

Anacapa: SBBG.

San Nicolas: SBBG.

Santa Catalina: Fosberg S4308 LAM!; Blakley 5530-A DS!;
RSA-POM.

San Clemente: Elmore s.n. Nov. 26, 1939 AHFH!; Blakley
6346 DS!; Piehl 62366 DS!; SBBG.

Stipa columbiana Macoun var. nelsoni (Scribn.) Hitchc.

San Nicolas: Trask 1 1 in Apr. 1897 US#340330!.

Stipa lepida Hitchc.

San Miguel: SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Hoffmann s.n. Jun. 14, 1930 LAM!; Fosberg
7610 LAM!, LA!; Brandegee 74 US!; DS; RSA-POM;
SBBG; SBM.

Anacapa: SBBG; SBM.

Santa Barbara: Dunkle 8109 LAM!.

Santa Catalina: Fosberg S4609 LAM!; Trask s.n. in Mar.

1901 NY!; Brandegee 59 US!; RSA-POM; SBM.

San Clemente: RSA-POM.

Guadalupe: Wiggins & Ernst 203 DS!, SD!; Moran 6754
RSA!; Wiggins & Ernst 199 DS!.

Stipa pulchra Hitchc.

San Miguel: Blakley 5838 DS!; SBBG; SBM.

Santa Rosa: RSA-POM; SBBG; SBM.

Santa Cruz: Wolf 2772 DS!; Hoffmann s.n. Mar. 23, 1929
CAS# 168484!; RSA-POM; SBBG; SBM.

Anacapa: SBBG; SBM.

San Nicolas: Trask 16 US!; RSA-POM; SBM.

Santa Barbara: Philbrick 68-83 SBBG!; Piehl 631631 CAS!.
Santa Catalina: Dunkle 1 780 AHFH!; Dunkle 1 786 AHFH!;

Knopf 201 US!; RSA-POM; SBM.

San Clemente: Trask 361 US!; Moran 570 LAM!, DS!;
Dunkle 7360 AHFH!; Abrams & Wiggins 381 CAS!,
DS!; RSA-POM.

Triticum aestivum L.

San Clemente: RSA-POM.

Triticum cylindricum (Host.) Ces., Pass. & Gib.

Santa Cruz: Barbe 1566 RSA!.

Vulpia bromoides (L.) S.F. Gray
San Miguel: Hoffmann s.n. Apr. 20, 1932 SBM# 11908!;
Hoffmann s.n. Apr. 20, 1932 SBM# 11926!; Hoffmann
s.n. Jun. 11, 1930 SBM#4991!.

Santa Rosa: Munz & Crow 11657 LA!; Hoffmann s.n. Apr.
16, 1929 SBM#6254!; Hoffmann s.n. Apr. 17, 1929
SBM#6256!; RSA-POM.

Santa Cruz: Hoffmann s.n. Jun. 29, 1930 LAM!, SBM!;
Clokey 4871 LAM!, NY!, US!; Abrams & Wiggins 101
DS!, NY!; SBBG; SBM.

Anacapa: Hoffmann s.n. Mar. 16, 1929 CAS# 168488!;

SBBG; SBM.

Santa Catalina: SBBG.

San Clemente: RSA-POM.

Guadalupe: RSA-POM.

Vulpia microstachys (Nutt.) Benth. var. pauciflora (Beal)
Lonard & Gould
San Miguel: SBM.

Anacapa: SBBG.

Santa Catalina: Fosberg S4543 LAM!; Fosberg S4728
LAM!, NY!, SBM!; Trask G-22 in Mar. 1 898 US!; RSA-
POM.

San Clemente: Trask 323 US!; RSA-POM; SBBG.
Guadalupe: Greene 42 in Apr. 1 885 US!; Moran 6658 SD!;
Wiggins & Ernst 1 14 DS!.

Vulpia myuros (L.) K.C. Gmelin var. hirsuta Hack.

San Miguel: Hoffmann s.n. Apr. 19, 1932 SBM# 1 1890!;
SBBG.

Santa Rosa: Hoffmann s.n. May 10, 1932 SBM#12159!;
Hoffmann s.n. Mar. 24, 1927 SBM!; Hoffmann s.n. Apr.
18, 1929 SBM#6255!; RSA-POM; SBBG.

Santa Cruz: Clokey 4862 LAM!; Dunkle 8640 LAM!; Bran-
degee s.n. in 1888 UC!; RSA-POM; SBBG; SBM.
Anacapa: [Hoffmann] s.n. Mar. 11, 1928 SBM#4092!;

SBBG; SBM.

San Nicolas: SBBG.

Santa Barbara: Dunkle 8140 LAM!, AHFH!; RSA-POM.
Santa Catalina: Trask G-19 in May 1898 US!; Fosberg
S4306 LAM!, NY!; Nuttall 98 NY!, US!; RSA-POM;
SBM.

90 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

San Clemente: Trask 360 NY!, US!; Dunkle 7257 LAM!,
AHFH!; Dunkle 7297 AHFH!; RSA-POM.

Guadalupe: Palmer 672 NY!; Palmer 673 US!; Moran
12387 SD!.

Vu/pia myuros (L.) K.C. Gmelin var. myuros
Santa Catalina: Brandegee 53 US!; Millspaugh 4667 US!;

Millspaugh 4679 US!; RSA-POM.

San Clemente: RSA-POM.

Vulpia octoflora (Walt.) Rydb. var. hirtella (Piper) Henr.
San Miguel: Hoffmann s.n. Apr. 20, 1932 SBM# 1 1898!;
SBBG.

Santa Rosa: Hoffmann s.n. Apr. 17, 1929 SBM#6262!;
RSA-POM; SBM.

Santa Cruz: Hoffmann s.n. Mar. 21, 1932 SBM# 11770!;

Hoffmann 256 SBM!; Abrams & Wiggins 46 DS!.
Anacapa: SBBG; SBM.

San Nicolas: Howell 8222 CAS!.

Santa Barbara: SBBG.

Santa Catalina: Brandegee 49 US!; Grant 3790 US!; Trask
s.n. in Mar. 1901 MO!, NY!, US!; RSA-POM.

San Clemente: RSA-POM; SBBG.

Guadalupe: Palmer 97 NY!; Palmer 637 NY!; Moran 6741
SD!; Howell 8309 CAS!, DS!.

Potamogetonaceae

Potamogeton crispus L.

Santa Catalina: RSA-POM.

Potamogeton pectinatus L.

Santa Rosa: RSA-POM.

Santa Cruz: Clokey 4854 LAM!, LA!; RSA-POM; SBM.
Santa Catalina: RSA-POM.

Ruppia martima L.

Santa Rosa: Hoffmann s.n. Jun. 13, 1930 DS!; Hoffmann
s.n. Apr. 28, 1930 CAS!; RSA-POM; SBM.

Santa Cruz: Daily 152 SCIR!.

Santa Catalina: Fosberg S4754 LAM!; Wolf 359 1 DS!; RSA-
POM; SBM.

San Clemente: RSA-POM.

Typhaceae

Typha domingensis Pers.

San Miguel: SBBG.

Santa Rosa: RSA-POM.

Santa Cruz: SBBG; SBM.

San Nicolas: Blakley 4158 SBBG!.

Santa Catalina: Trask s.n. in Mar. 1897 NY!.

Typha latifolia L.

San Nicolas: RSA-POM; SBBG.

Santa Catalina: Dunkle 2010 LAM!, AHFH!; Fosberg s.n.

LAM!; RSA-POM; SBBG.

San Clemente: Raven 18018 RSA!; SBBG.

Zosteraceae

Phyllospadix scouleri Hook.

Anacapa: RSA-POM.

San Nicolas: RSA-POM.

Santa Barbara: RSA-POM.

Santa Catalina: RSA-POM.

San Clemente: RSA-POM.

Phyllospadix torreyi Wats.

San Miguel: SBBG; SBM.

Santa Rosa: RSA-POM; SBM.

Santa Cruz: RSA-POM; SBM.

Anacapa: SBBG.

San Nicolas: RSA-POM.

Santa Barbara: Philbrick & Benedict B70-35 SBBG!.

Santa Catalina: Fosberg S4575 LAM!; Fosberg S4877
LAM!; Fosberg S4757 LAM!; RSA-POM.

San Clemente: RSA-POM.

Guadalupe: Moran 1 7422 RSA!, SD!; Moran 18156 RSA!.
Zostera marina L.

Santa Rosa: SBM.

Santa Cruz: Dunkle 8568 LAM!; Johnstone s.n. in Dec.
1928 USC!; SBBG; SBM.

Anacapa: Dunkle 7671 LAM!; RSA-POM; SBM.

Santa Catalina: Fosberg S4750 LAM!; RSA-POM.

Guadalupe: Hubbs 19290 LAM!; Moran 7844 SD!.

APPENDIX II. INDEX TO THE DISPOSITION
OF SYNONYMS, MISIDENTIFICATIONS, AND
TAX A INCERTAE SEDIS

It is the intent of this section to indicate the disposition in
this paper of taxa known to at least some earlier authors
under names not here accepted, not to provide a technical
synonymy of all insular taxa. Some of the names given here
are indeed synonymous. Many are based on earlier concepts
of certain taxa which were more inclusive or fragmented than
those currently accepted. Excluded, for some saving of space,
are all synonyms recognized in Munz and Keck (1959). It is
hoped that readers will find this section useful for locating
otherwise obscure reports and specimens upon which they
are based. There is a need to document more fully the elim-
ination of records from a flora to avoid interminable spec-
ulation as to their origin and fate.

The arrangement of this section is alphabetical by genus
and species. First, the combination in question is at the left
margin. In most cases, author citations for these names are
corrected from its citation, if it was in error. Second, just
below and indented is the name of the taxon under which
the material or record is included here. Third, again below
and indented from the above, a chronological listing of the
references using the combination in question. Each entry
consists of the authority, date of publication and an abbre-
viated notation of the insular occurrences to which the name
applies. In many cases the specimens, duplicates of the col-
lections, or the specimens upon which these reports most
probably are based have been located. In these cases the
specimens are cited just preceding the notation of the island.
In the case of literature citations, very often subsequent au-
thors simply repeat the citations of earlier workers without
seeking out the specimens upon which the record is based.

Occasionally it was not possible to reach a decision as to
the disposition of some reports. These are listed as incertae
sedis. In a few cases a reference documenting the placement
of certain taxa is noted following the accepted name used
here (e.g., see Baeria).

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 91

Abronia alba Eastw.

Abronia umbellata Lam. with introgression from A. ma-
ritima Nutt, ex Wats, omitted (Tillett, 1967).
Eastwood (1898) NI; Davidson and Moxley (1923) NI,
CL; Jepson (1925) NI; Hoffmann (1932a) MI, RO;
Munz ( 1 935) MI, RO, NI, CL; Eastwood ( 1 94 1 ) MI,
RO, NI; Gentry (1949) MI, RO, NI, CL; Dunkle
(1950) MI, RO, NI, CL.

Abronia alba Eastw. var. platyphylla (Standi.) Jeps.

Abronia umbellata Lam. with introgression from A. ma-
ritima Nutt, ex Wats, omitted (Tillett, 1967).
Hoffmann (1932a) RO; Eastwood (1941) RO.

Abronia alba Eastw. var. variabilis Jeps.

Abronia umbellata Lam. with introgression from A. ma-
ritima Nutt, ex Wats, omitted (Tillett, 1967).
Hoffmann (1932a) RO, CR.

Abronia insularis Standi.

Abronia umbellata Lam. with introgression from A. ma-
ritima Nutt, ex Wats, omitted (Tillett, 1967).
Eastwood (1941) CL.

Abronia minor Standi.

Abronia umbellata Lam. with introgression from A. lati-
folia Esch. omitted (Tillett, 1967).

Eastwood (1941) RO, CR.

Abronia neurophylla Standi.

Abronia umbellata Lam. with introgression from A. ma-
ritima Nutt, ex Wats, omitted (Tillett, 1967).
Eastwood (1941) NI.

Abronia umbellata Lam. ssp. alba (Eastw.) Munz

Abronia umbellata Lam. with introgression from A. ma-
ritima Nutt, ex Wats, omitted (Tillett, 1967).

Munz and Keck (1959) MI, RO, NI, CL.

Achillea borealis Bong.

Achillea millefolium L.

Philbrick (1972) BA.

Achillea lanulosa Nutt.

Achillea millefolium L.

Millspaugh and Nuttall (1923) CA; Eastwood (1941) MI,
RO, CR, AN, NI, BA, CA, CL; Raven (1963) CL;
Foreman (1967) NI.

Achillea millefolium L. lanulosa Piper
Achillea millefolium L.

Dunkle (1950) MI, RO, CR, AN, NI, BA, CA, CL.
Achyrachaena mollis Schauer

omitted as unsubstantiated on CA.

Brandegee (1890a) CA; Brandegee (1890b) CA; East-
wood (1941) CA.

Acrolasia gracilenta Rydb.

Mentzelia affinis Greene

Millspaugh and Nuttall (1923) CA.

Adenostoma fasciculatum H. & A. var. obtusifolium Wats.
Adenostoma fasciculatum H. & A. war. fasciculatum
Jepson (1909-1943) RO; Eastwood (1941) CR.
Adenostoma fasciculatum H. & A. var. prostratum Dunkle
Adenostoma fasciculatum H. & A. var. fasciculatum
Dunkle (1 94 1 ) based on Dunkle 8496 LAM! RO; Dunkle
(1950) RO.

Adiantum capillus-veneris L.

omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Munz (1935) CL.

Adiantum jordani C. Muell.

omitted as column transposition for CA.

Dunkle (1950) p. 293, BA.

Agoseris heterophylla Greene

omitted as unsubstantiated on NI.

Dunkle (1950) NI.

possibly Malacothrix saxatilis (Nutt.) T. & G.

Foreman (1967) based on Foreman 134 UC!, misiden-
tified, NI.

Agoseris heterophylla Greene

probably Agoseris grandiflora (Nutt.) Greene
Dunkle (1950) MI.

Agropyron repens (L.) Beauv.
incertae sedis

Greene (1887a) CR; Greene (1887b) MI; Brandegee
(1888) RO; Yates (1889) MI, RO, CR; Brandegee
(1890a) CA; Brandegee (1890b) MI, RO, CR, CA;
Eastwood (1941) MI, RO, CR, CA; Smith (1976)
MI.

Lolium perenne L. ssp. perenne

Foreman (1967) based on Foreman 111 UC!, misiden-
tified, NI.

Agrostis canina Bubani
Agrostis diegoensis Vasey

Brandegee (1890a) CA; Brandegee (1890b) CA.
Agrostis exarata Trin.

Agrostis diegoensis Vasey

Eastwood (1941) probably based on Abrams & Wiggins
370 DS!, US!, misidentified; also by Raven (1963)
CL.

Agrostis exarata Trin.

Polypogon interruptus HBK.

Brandegee (1890a) probably based on Brandegee s.n.
Mar. 13, 1890 UC!, misidentified, CA; Brandegee
(1890b) CA; Millspaugh and Nuttall (1923) CA;
Eastwood (1941) CA.

Agrostis foliosa Vasey
Agrostis diegoensis Vasey
Jepson (1909-1943) RO, CA.

Agrostis microphylla Steud.

Agrostis exarata Trin.

Hoffmann ( 1 932a) probably based on Hoffmann s.n. Aug.
7, 1930 SBM#10765!, misidentified, RO; Smith
(1976) RO.

Agrostis scouleri Trin.

Agrostis exarata Trin.

Brandegee (1888) RO; Yates (1889) RO; Brandegee
(1890b) RO; Eastwood (1941) RO, CR.

Agrostis vertici/lata Vill.

Agrostis semiverticillatus (Forsk.) Christensen

Brandegee (1888) RO; Yates (1889) RO; Brandegee
(1890b) RO; Millspaugh and Nuttall (1923) CA;
Hoffmann (1932a) CR; Eastwood (1941) RO, CR,
CA.

92 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Allium amplectens Torr. in T. & G.
incertae sedis
Eastwood (1941) CR.

Allium hyalinum Curran
Allium praecox Bdg.

Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890b) CR.

Allium peninsulare Lemmon
Allium praecox Bdg.

Jepson ( 1 909- 1 943) CA; Eastwood (1941) CA, CL; Munz
and Keck ( 1 959) CA, CL; Raven (1963) CL; Thome
(1967) CA; Wiggins (1980) CA, CL.

Allium serratum Wats.

Allium praecox Bdg.

Lyon (1886) CL; Brandegee (1890a) CA; Brandegee
( 1 890b) CA, CL; Millspaugh and Nuttall ( 1 923) CA.
Allocarya acanthocarpa Piper
incertae sedis
Wiggins (1980) GU.

Amaranthus blitoides Wats.

omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Eastwood (1941) CL.

Amaranthus graecizans L.

Amaranthus albus L.

Millspaugh and Nuttall (1923) CA; Hoffmann (1932a)
RO, CR; Eastwood (1941) MI, RO, CR, CA.

Ambrosia calif ornica Rydb.

Ambrosia psilostachya DC. var. californica (Rydb.) Blake
in Tides.

Eastwood (1941) CR, CA.

Ambrosia chamissonis Less.

omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Foreman (1967) BA.

Ambrosia psilostachya DC. var. californica (Rydb.) Blake in
Tides.

omitted as unsubstantiated on RO.

Smith (1976) RO.

Amsinckia catalinensis Suksd.

Amsinckia intermedia F. & M. (Jepson, 1909-1943)
Eastwood (1941) CA.

Amsinckia congesta Suks.

Amsinckia intermedia F. & M. (Jepson, 1909-1943)
Eastwood (1941) CA.

Amsinckia douglasiana A. DC.
incertae sedis
Clokey (1931) CR.

Amsinckia intermedia F. & M.

Millspaugh and Nuttall (1923) based in part on Mill-
spaugh 4616 F!, Millspaugh 4725 F!, Millspaugh
4796 F!, Millspaugh 4890 F!, Nuttall 46 F!, all mis-
identified, CA.

Amsinckia evermannii Suks.

Amsinckia intermedia F. & M. (Jepson, 1909-1943)
Eastwood (1941) CA, based on Evermann s.n. Mar. 25,
1918 CAS#26982!, CL.

Amsinckia insularis Suks.

Amsinckia intermedia F. & M. (Jepson, 1909-1943)
Eastwood (1941) CA, CL.

Amsinckia intermedia F. & M.

Amsinckia spectabilis F. & M. var. spectabilis

Macbride (1917) based on Trask 59 GH!, misidentified,
NI.

Amsinckia lycopsoides Lehm.
incertae sedis

Greene (1887a) CR; Brandegee (1890a) CA; Brandegee
(1890b) CR, CA; Davidson (1896) CA.

Amsinckia spectabilis F. & M. var. spectabilis

Greene (1887b) probably based on Greene s.n. in Sep.
1886 ND-G#042845!, misidentified, MI; Yates
(1889) MI; Brandegee (1890b) MI.

Amsinckia maritima Eastw.
incertae sedis
Eastwood (1941), MI, CL.

Amsinckia spectabilis F. & M. var. spectabilis

Eastwood (1898) based on Trask 59 GH!, misidentified,
NI; Eastwood (1941) NI.

Amsinckia microsperma Suks.

Amsinckia spectabilis F. & M. var. spectabilis (Jepson,
1909-1943)

Eastwood (1941) CA.

Amsinckia nesophila Suks.
incertae sedis
Eastwood (1941) CA.

Amsinckia sanctinicolai Eastw.

Amsinckia spectabilis F & M. var. nicolai (Jeps.) Jtn. ex
Munz

Jepson (1909-1943) NI.

Amsinckia spectabilis F. & M.

Amsinckia intermedia F. & M.

Brandegee (1890b, in footnote) CA.

Amsinckia St. Nicolai Eastw.

omitted as unsubstantiated on CR.

Eastwood (1941) CR.

Amsinckia swainiae Suks.

Amsinckia intermedia F. & M. (Jepson, 1909-1943)
Eastwood (1941) CR, CA.

Amsinckia tessellata Gray
incertae sedis

Brandegee (1890a) CA; Davidson (1896) CA.
Amsinckia vernicosa H. & A.

Amsinckia menziesii (Lehm.) Nels. & Macbr.

Watson (1876) based on Palmer 69 GH!, MO!, NY!,
misidentified, GU; Eastwood (1929) GU; Dunkle
(1950) GU.

Anaphalis margaritacea (L.) Benth. & Hook.

omitted as probable error; also omitted by Hall (1907).
Brandegee (1890a) CA; Brandegee (1890b) CA; David-
son (1896) CA; Eastwood (1941) CA.

Andropogon barbinodis Lag.

Bothriochloa barbinodis (Lag.) Herter
Abrams (1917) CA; Thome (1967) CA.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 93

Andropogon saccharoides Swartz

Bothriochloa barbinodis (Lag.) Herter

Davidson ( 1 894) CA; Millspaugh and Nuttall ( 1 923) CA;
Eastwood (1941) RO, CA.

Antennaria margaritacea Benth. nom. nud. in lit.
omitted as probable error.

Brandegee (1890a) CA.

Aphyllon fasciculata (Nutt.) T. & G. ex Gray
Orobanche fasciculata Nutt.

Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890b) CR.

Aplopappus canus (Gray) Blake
Haplopappus canus (Gray) Blake

Munz (1935) CL, GU; Dunkle (1950) CL, GU.
Haplopappus detonsus (Greene) Raven

Munz ( 1 93 5) RO, CR; Dunkle (1942) AN; Dunkle (1950)
RO, CR, AN.

Aplopappus ericoides (Less.) H. & A.

Haplopappus ericoides (Less.) H. & A. ssp. ericoides
Greene ( 1 887b) MI; Yates ( 1 889) MI; Brandegee ( 1 890b)
MI; Munz (1935) MI.

Aplopappus squarrosus H. & A.

Haplopappus squarrosus H. & A. ssp. grindelioides (DC.)
Keck

Greene (1887a) CR; Yates (1889) CR; Brandegee ( 1 890b)
CR.

Aplopappus venetus (HBK.) Blake var. furfuraceus (Greene)
Munz

omitted as column transposition for CA.

Dunkle (1950, p. 293) BA.
omitted as unsubstantiated on CR.

Dunkle (1950) CR.

Haplopappus venetus (HBK.) Blake ssp. furfuraceus
(Greene) Hall

Munz (1935) CA, CL; Dunkle (1950) CA, CL.
Aplopappus venetus (HBK.) Blake var. sedoides (Greene) Munz
Haplopappus venetus (HBK.) Blake ssp. sedoides (Greene)
Munz

Munz (1935) RO, CR; Dunkle (1950) RO, CR.
Aplopappus venetus (HBK.) Blake var. vernonioides (Nutt)
Munz

omitted as column transposition for CA.

Dunkle (1950, p. 293) BA.

Haplopappus venetus (HBK.) Blake ssp. vernonioides (Nutt.)
Hall

Dunkle (1942) AN; Dunkle (1950) MI, RO, CR, NI, CL.
Arabis arcuata Gray

Arabis hoffmannii (Munz) Roll.

Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890b) CR.

Arctostaphylos sp.
incertae sedis

Greene (1885) GU; Eastwood (1929) GU.
Arctostaphylos andersonii Gray var. viridissima (Eastw.) Jeps.
Arctostaphylos catalinae P.V. Wells
Jepson (1909-1943) CA.

Arctostaphylos confertiflora Eastw.

Jepson (1909-1943) RO.

Arctostaphylos viridissima (Eastw.) McMinn
Jepson (1909-1943) CR.

Arctostaphylos Crustacea Eastw.

probably Arctostaphylos tomentosa (Pursh) Lindl. ssp. in-
sulicola P.V. Wells.

Munz and Keck (1959) RO, CR.

Arctostaphylos divers if oli a Parry

Comarostaphylis diversifolia (Parry) Greene ssp. planifolia
(Jeps.) Wallace ex Thome

Brandegee (1888) RO; Yates (1889) RO, CR; Ford ( 1 890)
CR; Brandegee ( 1 890a) CA; Brandegee ( 1 890b) RO,
CR, CA; Davidson (1896) CA; Trask (1899) CA;
Munz (1935) RO, CR, CA; Dunkle (1950) RO, CR,
CA.

Arctostaphylos glandulosa Eastw.

Arctostaphylos tomentosa (Pursh) Lindl. ssp. subcordata
(Eastw.) P.V. Wells.

Jepson (1909-1943), in references CR.

Arctostaphylos insularis Greene
incertae sedis
Dunkle (1950) CR.

Arctostaphylos catalinae P.V. Wells

Davidson (1896) CA; Munz (1935) CA; Dunkle (1950)
CA.

Arctostaphylos confertiflora Eastw.

Munz (1935) based on Munz & Crow 1 1587 POM!,
misidentified, RO; Eastwood (1941) RO; Dunkle
(1950) RO; Smith (1976) RO.

Arctostaphylos pechoensis Dudley var. viridissima Eastw.
Arctostaphylos catalinae P.V. Wells
Munz and Keck (1959) CA.

Arctostaphylos viridissima (Eastw.) McMinn

Abrams and Ferris (1923-1960) CR; Munz (1935) CR;
Eastwood (1941) CR; Munz and Keck (1959) CR.
Arctostaphylos pungens HBK.

Arctostaphylos catalinae P.V. Wells
Brandegee (1890a) CA.

Arctostaphylos stanfordiana Parry
incertae sedis
Yates (1889) CR.

Arctostaphylos subcordata Eastw.

Arctostaphylos catalinae P.V. Wells
Thome (1967) CA.

Arctostaphylos tomentosa (Pursh) Lindl. ssp. subcordata
(Eastw.) P.V. Wells

Abrams and Ferris (1923-1960) CR; Munz (1935) RO,
CR; Eastwood (1941) CR; Dunkle (1950) RO, CR;
Munz and Keck (1959) CR.

Arctostaphylos subcordata Eastw. var. confertiflora (Eastw.)
Munz

Arctostaphylos confertiflora Eastw.

Munz and Keck (1959) RO.

Arctostaphylos tomentosa (Pursh) Lindl.
incertae sedis

Greene ( 1 887a) CR; Brandegee ( 1 888) RO; Yates ( 1 889)
RO, CR; Brandegee ( 1 890b) RO, CR; Jepson ( 1 909-
1 943) CR; Hoffmann (1932b) CR; Munz ( 1 935) CR;
Eastwood (1941) RO, CR; Dunkle (1950) RO, CR.

94 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Arctostaphylos catalinae P.V. Wells

Lyon (1886) CA; Brandegee (1890b) CA; Munz (1935)
CA; Eastwood (1941) CA; Dunkle (1950) CA.
Arctostaphylos tomentosa (Pursh) Lindl. var. hispida Hook,
nom. nud. pro. syn.

Arctostaphylos tomentosa (Pursh) Lindl. ssp. insulicola P.V.
Wells

Hoffmann (1932b) based on Hoffmann s.n. Jun. 29, 1930
SBM#5736!, annotated by Eastwood but no such
combination found, misidentihed, CR.

Artemisia californica Less. f. erecto Dunkle nom. nud. pro.
syn.

Artemisia californica Less.

Dunkle (1950) MI, RO, CR, CA.

Artemisia californica Less. f. flexila Dunkle nom. nud. pro.
syn.

Artemisia californica Less.

Dunkle (1950) MI, RO, CR, CA.

Artemisia californica Less. var. insularis (Rydb.) Munz
Artemisia californica Less.

Eastwood (1941) MI, RO, CR, AN, CA; Dunkle (1950)
AN.

Artemisia nesiotica Raven

Abrams and Ferris (1923-1960) NI, CL; Howell (1935)
NI; Munz (1935) NI, CL; Eastwood (1941) NI, BA,
CL; Dunkle (1942) BA; Dunkle (1950) NI, BA, CL.
Artemisia ludoviciana Nutt.

Artemisia douglasiana Bess, in Hook.

Greene (1887a) CR; Brandegee ( 1 888) RO; Yates ( 1 889)
RO, CR; Ford (1890) CR; Eastwood (1941) RO,
CR.

Artemisia vulgaris L.

Artemisia douglasiana Bess, in Hook.

Brandegee (1890b) ref. as same as A. ludoviciana of pre-
vious lists, RO, CR, CA; Davidson (1896) CA;
Millspaugh and Nuttall (1923) CA.

Aspidium sp.

Athyrium felix-femina (L.) Roth var. sitchensis Rupr. (ref.
Brandegee, 1890b).

Greene (1887a) CR.

Aspidium aculeatum Swartz
Dryopteris arguta (Kaulf.) Watt

Lyon (1886) CA; Brandegee (1890b) CA.

Aspidium munitum Kaulf.

Polystichum munitum (Kaulf.) Presl ssp. solitarium Maxon
Watson (1876) GU.

Aspidium rigidum Swartz

Dryopteris arguta (Kaulf.) Watt
Greene (1887a) CR; Brandegee (1888) RO; Brandegee
(1890a) CA; Brandegee ( 1 890b) RO, CR, CA; Trask
(1899) CA.

Asplenium filix-foemina Bemh. [sic]

Athyrium felix-femina (L.) Roth. var. sitchensis Rupr.
Brandegee (1890b) CR.

Aster foliaceus Lindl.

Aster chilensis Nees var. chilensis

Brandegee (1888) RO; Yates (1889) RO; Brandegee
(1890b) RO.

Astragalus antiselli Gray

Astragalus trichopodus (Nutt.) Gray ssp. trichopodus
Lyon (1886) CA; Brandegee (1890b) CA.

Astragalus didymocarpus H. & A.

omitted as unsubstantiated on NI; also omitted by Barneby
(1964).

Eastwood (1898) NI.

Astragalus doug/asii Gray
Astragalus curtipes Gray

Munz (1935) MI; Dunkle (1950) MI.

Astragalus miguelensis Greene

Dunkle (1942) probably based on Elmore 252 AHFH!,
misidentihed, AN; Dunkle (1950) AN.

Astragalus fastidiosus Greene
Astragalus trichopodus (Nutt.) Gray ssp. leucopsis (T. &
G.) Thorne
Eastwood (1941) CA.

Astragalus leucopsis (T. & G.) Torr.

omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Eastwood (1941) BA; Dunkle ( 1 942) BA; Gentry ( 1 949)
BA; Dunkle (1950) BA.

Astragalus curtipes Gray

Brandegee (1890b) probably based on Greene (1887b),
misidentihed, MI, see also Barneby ( 1 964) reference
for A. leucopsis (T. & G.)Torr. var. brachypus Greene
Astragalus miguelensis Greene

Gentry (1949) based on Elmore 319 AHFH!, misiden-
tihed, MI, based on Elmore 252 AHFH!, misiden-
tihed, AN; Dunkle (1950) MI, AN in part.
Astragalus trichopodus (Nutt.) Gray ssp. leucopsis (T. &
G.) Thome

Lyon ( 1 886) CA; Greene ( 1 887a) CR; Brandegee (1888)
RO; Yates (1889) CR; Brandegee (1890b) RO, CR,
CA; Trask ( 1 899) CA; Jepson ( 1 909-1943) CR, CA;
Munz (1935) CR, CA; Eastwood (1941) CA; Dunkle
(1942) AN; Gentry (1949) CA; Dunkle (1950) RO,
CR, AN in part; Munz and Keck (1959) CR, CA;
Thome (1967) CA; Thome (1969) CA.

Astragalus leucopsis (T. & G.) Torr. var. brachypus Greene
probably Astragalus trichopodus (Nutt.) Gray ssp. leucopsis
(T. & G.) Thome.

Dunkle (1950) CR.

Astragalus nevinii Gray

omitted as unsubstantiated on CA; also omitted by Bar-
neby (1964).

Davidson and Moxley (1923) CA; Eastwood (1941) CA;
Gentry (1949) CA.

Astragalus miguelensis Greene (Barneby, 1964, p. 464)
Jepson (1909-1943) based on Hemphill [at UC], mis-
identihed, AN; Eastwood ( 1 941) AN; Gentry (1949)
AN.

Astragalus traskiae Eastw.

Jepson (1909-1943) BA; Eastwood (1941) BA; Dunkle
(1942) BA; Gentry (1949) BA; Dunkle (1950) BA.

Astragalus robeartsii Eastw.
incertae sedis
Trask (1904) CL.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 95

Astragalus traskiae Eastw.

omitted as column transposition for CA.

Dunkle (1950, p. 293) BA.

Astragalus miguelensis Greene (Bameby, 1964, p. 462)
Abrams and Ferris (1923-1960) AN.

Astragalus nevinii Gray (Bameby, 1964, p. 462)
Eastwood (1941) CL; Dunkle (1950) CL.

Astragalus trichopodus (Nutt.) Gray var. gaviotus (Elmer)
Jeps.

Astragalus trichopodus (Nutt.) Gray ssp. trichopodus
Jepson (1925) CA.

Astragalus trichopodus (Nutt.) Gray var. lonchus (Jones) Bar-
neby

Astragalus trichopodus (Nutt.) Gray ssp. leucopsis (T. &
G.) Thome

Bameby (1964) CR, AN, CA.

Atriplex bracteosa (Dur. & Hilg.) Wats.

Atriplex serenana A. Nels. var. serenana
Hoffmann (1932a) RO.

Atriplex coulteri (Moq.) D. Dietr.

omitted as column transposition for CA.

Dunkle (1950, p. 293) BA.

Atriplex microcarpa (Benth.) D. Dietr.

Atriplex pacifica Nels.

Greene (1887a) based on Greene s.n. in Jul.-Aug. 1886
ND-G#0 15274!, US!, misidentified, CR.

Atriplex pacifica Nels.

omitted as column transposition for CA.

Dunkle (1950, p. 293) BA.

Atriplex coulteri (Moq.) D. Dietr.

Munz (1935) probably based on Hoffmann s.n. Jun. 13,
1930 POM!, misidentified, RO; Eastwood (1941)
RO; Dunkle (1950) RO; Munz and Keck (1959) RO;
Smith (1976) RO.

Atriplex palmeri Wats.

Atriplex barclayana (Benth.) D. Dietr. ssp. palmeri ( Wats.)
Hall & Clem.

Watson (1876) GU; Greene (1885) GU; Eastwood (1929)
GU.

Atriplex rosea L.

Aphanisma blitoides Nutt, ex Moq. in DC.

Dunkle (1942) based on Dunkle 7459 AHFH!, misiden-
tified, BA; Dunkle (1950) BA.

Atriplex rosei Standi.

Atriplex barclayana (Benth.) D. Dietr. ssp. dilitata (Greene)
Hall & Clem.

Eastwood (1929) GU.

Atriplex serenana A. Nels. var. davidsonii (Standi.) Munz
Atriplex serenana A. Nels. var. serenana

Eastwood (1941) RO, CR; Thome (1967) based on East-
wood 6529 US!, misidentified, CA; Smith (1976)
RO.

Atriplex watsonii A. Nels. in Abrams
omitted as column transposition for CA.

Dunkle (1950, p. 293) BA.

Audibertia nivea Benth.
incertae sedis

Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890b) CR; Ford (1890) CR.

Audibertia palmeri Gray
Salvia mellifera Greene

Lyon (1886) CA; Greene (1887a) CR; Yates (1889) CR;
Ford (1890) CR; Brandegee (1890b) CR, CA.
Baccharis viminea DC.

Baccharis glutinosa Pers.

Greene (1887a) CR; Yates (1889) CR; Ford (1890) CR;
Brandegee ( 1 890a) CA; Brandegee ( 1 890b) CR, CA;
Millspaugh and Nuttall (1923) CA; Eastwood (1941)
CR, CA; Dunkle (1942) AN; Munz and Keck (1959)
Channel Islands; Raven (1963) CL.

Baeria aristata Cov.

Lasthenia coronaria (Nutt.) Omduff
Dunkle (1950) GU.

Baeria chrysostoma F. & M.

Lasthenia californica DC. ex Lindl. (Johnson and Omduff,
1978)

Millspaugh and Nuttall (1923) CA.

Baeria chrysostoma F. & M. var. gracilis Hall
Lasthenia californica DC. ex Lindl. (Johnson and Omduff,
1978).

Munz (1935) on the islands; Dunkle (1942) AN; Gentry
(1949) CR, CL; Dunkle (1950) MI, RO, CR, AN,
NI, BA, CA, CL.

Baeria chrysostoma F. & M. ssp. hirsutula (Greene) Ferris
Lasthenia californica DC. ex Lindl. (Johnson and Omduff,
1978).

Munz and Keck (1959) Channel Islands.

Baeria chrysostoma F. & M. var. palmeri (Gray) J.T. Howell
Lasthenia californica DC. ex Lindl. (Johnson and Omduff,
1978).

Howell (1942) GU.

Baeria coronaria (Nutt.) Gray

Lasthenia coronaria (Nutt.) Omduff

Brandegee (1900) GU; Eastwood (1929) GU.

Baeria gracilis Gray

Lasthenia californica DC. ex Lindl. (Johnson and Omduff,
1978)

Brandegee (1888) RO, CR; Yates (1889) RO, CR, AN;
Brandegee ( 1 890a) CA; Brandegee ( 1890b) RO, CR,
CA; Eastwood (1898) NI; Brandegee (1900) GU;
Eastwood (1929) GU.

Baeria hirsutula Greene

Lasthenia californica DC. ex Lindl. (Johnson and Omduff,
1978).

Munz (1935) MI, RO, CR; Eastwood (1941) MI, RO,
CR, CA; Dunkle ( 1 942) BA; Dunkle (1950) MI, RO,
CR, NI, BA, CA.

Baeria macrantha (Gray) Gray
Lasthenia californica DC. ex Lindl. (Johnson and Omduff,
1978).

Munz and Keck (1959) MI, RO.

96 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Baeria palmeri Gray

Lasthenia californica DC. ex Lindl. (Johnson and Ornduff,
1978).

Watson (1876) GU; Greene (1885) GU; Vasey and Rose
(1890) GU; Eastwood (1929) GU.

Baeria palmeri Gray var. Clementina Gray
Lasthenia californica DC. ex Lindl. (Johnson and Ornduff,
1978).

Lyon ( 1 886) CA, CL; Greene (1887a) CR; Greene ( 1887b)
MI; Brandegee (1888) RO; Yates (1889) RO, CR;
Brandegee ( 1 890b) MI, RO, CR, CA, CL; Davidson
( 1 896) CA, CL; Howell (1935) NI; Eastwood (1941)
MI, RO, CR, NI, CA, CL.

Bahia lanata Nutt. var.

Eriophyllum lanatum (Pursh) Forbes var. grandiflorum
(Gray) Jeps.

Watson (1876) probably based on Palmer s.n. in 1875
GH!, GU; Eastwood (1929) GU.

Berula angustifolia Koch
Berula erecta (Huds.) Cov.

Greene (1887a) CR; Greene (1887b) MI; Yates (1889)
MI, CR; Brandegee (1890b) MI, CR; Eastwood
(1941) MI, CR.

Bigelovia veneta (HBK.) Gray
Haplopappus venetus (HBK.) Blake ssp. vernonioides ( Nutt.)
Hall

Lyon (1886) CA; Greene (1887a) CR; Greene (1887b)
MI; Brandegee (1888) RO; Yates (1889) MI, RO,
CR; Ford (1890) CR; Brandegee (1890b) MI, RO,
CR, CA.

Blepharipappus platyglossus (F. & M.) Greene
Layia platyglossa (F. & M.) Gray ssp. campestris Keck

Millspaugh and Nuttall (1923) CA.

Bloomeria aurea Kell.

Bloomeria crocea (Torr.) Cov. var. crocea

Greene (1887a) CR; Brandegee ( 1 888) RO; Yates (1889)
RO, CR; Brandegee ( 1 890a) CA; Brandegee ( 1 890b)
RO, CR, CA.

Brahea edulis Wendl.

Erythea edulis (Wendl.) Wats.

Watson (1876) GU.

Brassica campestris L.

Brassica rapa L. ssp. sylvestris (L.) Janchen

Greene (1 885) GU; Greene (1887b) MI; Brandegee (1888)
CR; Yates (1889) MI, CR; Brandegee (1890a) CA;
Brandegee (1890b) MI, CR, CA; Millspaugh and
Nuttall ( 1 923) CA; Eastwood ( 1 929) GU; Eastwood
(1941) MI, CR, CA; Smith (1976) RO, CR.

Brickellia californica (T. & G.) Gray
omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Dunkle (1950) CL.

Brodiaea sp.
incertae sedis

Howell (1942) GU.

Brodiaea capital a Benth.

Dichelostemma pulchellum (Salisb.) Heller

Greene (1885) GU; Lyon (1886) CL; Brandegee (1888)
RO; Vasey and Rose ( 1 890) GU; Brandegee ( 1 890a)
CA; Brandegee (1890b) MI, RO, CR, CA, CL; East-
wood ( 1 898) NI; Trask ( 1 899) CA; Munz ( 1 935) on
the islands; Howell (1935) NI; Eastwood (1941) MI,
RO, CR, NI, CA, CL; Dunkle (1942) AN; Gentry
(1949) CL; Dunkle (1950) MI, RO, CR, AN, NI,
BA, CA, CL, GU.

Brodiaea Clementina (Hoov.) Munz
Triteleia Clementina Hoov.

Munz and Keck (1959) CL.

Brodiaea flifolia Wats.

Brodiaea kinkiensis Niehaus

Munz and Keck (1959) CL.

Brodiaea insularis Greene

Dichelostemma pulchellum Heller

Greene (1887a) CR; Greene (1887b) MI; Yates (1889)
MI, CR; Eastwood (1929) GU.

Brodiaea laxa (Benth.) Wats.

Triteleia Clementina Hoov.

Munz (1935) CL; Dunkle (1950) CL.

Brodiaea lugens Greene

Triteleia guadalupensis Lenz

Brandegee (1900) GU; Eastwood (1929) GU; Munz and
Keck (1959) GU; Raven (1963) GU.

Brodiaea minor (Benth.) Wats.

Brodiaea jolonensis Eastw.

Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890a) CA; Brandegee (1890b) CR, CA; Trask
(1899) CA.

Brodiaea pulchella (Salisb.) Greene

Dichelostemma pulchellum (Salisb.) Heller

Foreman (1967) NI.

Brodiaea synandra (Heller) Jeps.

Brodiaea jolonensis Eastw.

Hoffmann (1932a) RO; Munz (1935) RO; Eastwood
(1941) RO, CR, CA.

Bromus sp.
incertae sedis

Greene (1887a) CR; Greene (1887b) MI.

Bromus carinatus H. & A.

Bromus maritimus (Piper) Hitchc.

Thome (1969) based on Fosberg S4484 (LAM!, mis-
identified, CA.

Bromus carinatus H. & A. var. hookerianus (Thurb. in Torr.)

Shear

Bromus carinatus H. & A.

Eastwood (1941) MI, RO, CR, NI, CA, CL; Dunkle
(1942) AN; Dunkle (1950) MI, RO, CR, AN, NI,
CA, CL.

Bromus ciliatus L.

possibly Bromus vulgaris (Hook.) Shear.

Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890a) CA; Brandegee (1890b) CR, CA.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 97

Bromus hookerianus Thurb. in Torr.

Bromus carinatus H. & A.

Brandegee (1888) RO, CR; Yates (1889) RO, CR; Bran-
degee ( 1 890a) CA; Brandegee ( 1 890b) RO, CR, CA,
CL; Eastwood (1898) NI, ref. to MI, RO, CR, CA,
CL.

Bromus laevipes Shear
omitted as unsubstantiated on AN.

Dunkle (1950) AN.

Bromus pseudolaevipes Wagnon
Hoffmann (1932a) probably based on [Holfmann] s.n. Jun.
13, 1930 SBM#10180!, misidentified, RO; Eastwood
(1941) RO; Holfmann (1932a) probably based on
Hoffmann s.n. Jun. 29, 1930 SBM#10132!, misiden-
tified, CR; Eastwood (1941) CR.

Bromus marginatus Nees in Steud.

Bromus carinatus H. & A.

Hoffmann (1932a) MI, RO, CR, AN; Eastwood (1941)
MI, RO, CR; Dunkle (1950) MI, RO, CR, AN, BA;
Smith (1976) MI, RO, CR, AN.

Bromus maritimus (Piper) Hitchc.

Thome (1967) based on Fosberg S4484 LAM!, mis-
identified, CA.

Bromus maximus Desf.

Bromus diandrus Roth.

McClatchie (1894) CA; Greene (1885) GU.

Bromus orcuttianus Vasey

Bromus pseudolaevipes Wagnon

Jepson (1909-1943) CA; Davidson and Moxley (1923)
CA; Millspaugh and Nuttall (1923) CA; Abrams
(1917) CA; Eastwood (1941) CA.

Bromus rigidus Roth, ex Reichenb.

Bromus diandrus Roth.

Millspaugh and Nuttall (1923) CA; Clokey (1931) CR;
Hoffmann (1932a) RO; Eastwood (1941) MI, RO,
CA; Dunkle ( 1 942) AN, BA; Dunkle (1950) MI, RO,
BA, CA.

Bromus rigidus Roth ex Reichenb. var. gussonei (Pari.) Coss.

& Dur.

Bromus diandrus Roth

Millspaugh and Nuttall (1923) CA; Hoffmann (1932a)
MI, RO, CR; Eastwood (1941) MI, RO, CR, NI,
CA.

Bromus sterilis L.
incertae sedis

Dunkle (1950) GU.

Bromus diandrus Roth.

Eastwood (1929) based in part on Mason 1543 GH!,
misidentified, GU.

Bromus rubens L.

Dunkle (1942) possibly based on Bond 373 SBM!, mis-
identified, BA; also denied by Philbrick (1972).
Bromus tectorum L.

Watson (1876) based on Palmer 99 (in part) NY!.
Bromus trinii Desv. in C. Gay

Watson (1876) based on Palmer 99 (in part) NY!.

Bromus subvelutinus Shear
Bromus carinatus H. & A.

Hoffmann ( 1 932a) based on Hoffmann s.n. Apr. 18,1 929
SBM#7649!, misidentified, RO; Eastwood (1941)
RO.

Bromus pseudolaevipes Wagnon

Millspaugh and Nuttall (1923) CA; Eastwood (1941) CA.
Bromus tectorum L.

Bromus sterilis L.

Millspaugh and Nuttall (1923) based in part on Nuttall
95 F!, Nuttall 562 F!, Nuttall 1219 F!, all misiden-
tified, CA; Eastwood (1941) CA.

Bromus virens Buckl.

Bromus carinatus H. & A.

Eastwood (1898) NI.

Bromus vulgaris (Hook.) Shear
Bromus trinii Desv. in C. Gay

Dunkle (1942) based on Dunkle 7242 AHFH!, misiden-
tified, BA; Dunkle (1950) BA.

Buda marina (L.) Dumort.

Spergularia marina (L.) Griesb.

Davidson (1896) CA.

Cakile edentula (Bigel.) Hook. var. californica Fem.

Cakile edentula (Bigel.) Hook. var. edentula
Hoffmann ( 1 932b) MI, RO; Munz (1935) MI; Munz and
Keck (1959) Channel Islands.

Calais linearifolia DC.

Microseris linearifolia (DC.) Sch.-Bip.

Greene (1887a) CR; Yates (1889) CR.

Calamintha palmeri Gray
Satureja palmeri (Gray) Briq.

Watson (1876) based on Palmer 66 NY!, GU; Greene
(1885) GU; Vasey and Rose (1890) GU; Eastwood
(1929) GU.

Calandrinia caulescens HBK.

Calandrinia ciliata (R. & P.) DC. var. menziesii (Hook.)
Macbr.

Hoffmann (1932a) MI, RO; Eastwood (1941) MI, RO,
CR, CA.

Calandrinia menziesii (Hook.) T. & G.

Calandrinia ciliata (R. & P.) DC. var. menziesii (Hook.)
Macbr.

Watson (1876) GU; Greene (1885) GU; Greene (1887a)
CR; Yates (1889) CR; Millspaugh and Nuttall (1923)
CA; Eastwood (1941) RO; Howell (1942) GU.
Calandrinia menziesii (Hook.) T. & G. var. caulescens (HBK.)
Gray

Calandrinia ciliata (R. & P.) DC. var. menziesii (Hook.)
Macbr.

Vasey and Rose (1890) GU.

Calochortus sp.
incertae sedis
Greene (1887a) CR.

Calochortus albus Dougl. ex Benth. var. rubellus Greene
Calochortus albus Dougl. ex Benth. var. albus
Clokey (1931) CR; Eastwood (1941) CR; Dunkle (1950)
CR; Smith (1976) CR.

98 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Calochortus kennedyi Porter
Calochortus cata/inae Wats.

Lyon (1886) CA.

Calochortus palmeri Wats.

Calochortus catalinae Wats.

Trask (1899) CA; Brandegee (1890b) CA.

Calochortus venustus Benth.

Calochortus catalinae Wats.

Brandegee (1888) CR; Yates (1889) CR.

Calystegia macrostegia (Greene) Brummitt ssp. cyclostegia
(House) Brummitt

omitted as unsubstantiated on CR; also omitted by Brum-
mitt pers. comm.

Smith (1976) CR.

Calystegia macrostegia (Greene) Brummitt ssp. macrostegia
Calystegia macrostegia (Greene) Brummitt ssp. amplissi-
ma Brummitt

Foreman (1967) NI; Philbrick (1972) Nl, BA, CL.
Camissonia guadalupensis (Wats.) Raven ssp. guadalupensis
Camissonia guadalupensis (Wats.) Raven ssp. Clementina
(Raven) Raven
Wiggins (1980) CL.

Caprifolium hispidulum Gray var. californicum Greene
Lonicera hispidula (Lindl.) Dougl. ex T. & G. var. vaci/lans
Gray

Davidson (1896) CA.

Capriola dactylon (L.) Kuntze
Cynodon dactylon (L.) Pers.

Millspaugh and Nuttall (1923) CA.

Capsella divaricata Walp.

Capsella bursa-pastoris (L.) Medic.

Greene (1887a) CR; Greene (1887b) MI; Brandegee
(1888) RO; Yates (1889) MI, RO, CR; Brandegee
(1890b) MI, RO, CR.

Cardamine integrifolia (Nutt.) Greene
Cardamine californica Greene
Greene (1887a) CR; Yates (1889) CR.

Cardamine paucisecta Benth.

Cardamine californica Greene
Brandegee (1890b) CR; Millspaugh and Nuttall (1923)
CA.

Carex sp.
incertae sedis

Greene (1887a) CR; Brandegee (1890a) CA.

Carex abrupt a Mkze.

Carex montereyensis Mkze.

Clokey (1931) based on Clokey 4874 LAM!, misiden-
tified, CR; Eastwood (1941) CR; Munz (1974) CR;
Smith (1976) CR.

Carex angustata Boott
incertae sedis

Greene (1887a) CR; Yates (1889) CR; Brandegee (1890b)
CR; Eastwood (1941) CR.

Carex douglasii Boott
incertae sedis

Brandegee (1888) RO; Yates (1889) RO; Brandegee
(1890b) RO; Eastwood (1941) RO, CR.

Carex glabra Boott
incertae sedis
Eastwood (1941) CR.

Carex praegracilis W. Boott
Carex tumulicola Mkze.

Munz (1935) probably based on Munz 6737 POM!, mis-
identified, CL.

Carex tumulicola Mkze.

Carex praegracilis W. Boott
Thome (1967) based on Thome 35902 RSA!, misiden-
tified, CA; Munz (1974) RO, CR.

Carpobrotus chilensis (Mol.) N.E. Br.

Carpobrotus aequilaterus (Haw.) N.E. Br.

Raven (1963) CL.

Castilleja affinis H. & A. var. contentiosa (Macbr.) Bacig.
Castilleja affinis H. & A. ssp. affinis
Smith (1976) RO, AN?; omitted since not recognized by
L. Heckard (pers. comm.).

Castilleja affinis H. & A. ssp. insularis (Eastw.) Munz
Castilleja affinis H. & A. ssp. affinis
Munz and Keck (1959) CR; Smith (1976) CR.
Castilleja anacapensis Dunkle

Castilleja affinis H. & A. ssp. affinis
Dunkle (1942) based on Dunkle 7639 AHFH!, LAM!,
and Dunkle 766 1 AHFH!, LAM!, AN; Gentry ( 1 949)
AN; Dunkle (1950) AN.

Castilleja foliolosa H. & A.
omitted as unsubstantiated on RO and CR.

Eastwood (1941) CR; Dunkle (1950) RO, CR.
Castilleja guadalupensis Bdg.

Watson (1876) GU; Eastwood (1 929) GU; Dunkle(1950)
GU.

Castilleja hololeuca Greene
omitted as unsubstantiated on CA and CL.

Jepson (1925) CA, CL.

Castilleja hololeuca Greene var. grisea (Dunkle) Munz
Castilleja grisea Dunkle
Munz and Keck (1959) CL.

Castilleja latifolia H. & A.
omitted as unsubstantiated on CR.

Gentry (1949) CR.

Castilleja mollis Penn.

Hoffmann (1932b) RO; Munz (1935) RO; Eastwood
(1941) RO; Gentry (1949) RO; Dunkle (1950) RO.
Castilleja latifolia H. & A. ssp. insularis Eastw.

Castilleja affinis H. & A. ssp. affinis
Eastwood (1941) CR.

Castilleja parviflora Bong.

Castilleja affinis H. & A. ssp. affinis
Lyon ( 1 886) CA; Brandegee ( 1 888) RO, CR; Yates ( 1 889)
RO, CR; Brandegee (1890b) RO, CR, CA.
Castilleja parviflora Bong. var. californica (Abrams) Zeile.
Castilleja affinis H. & A. ssp. affinis
Hoffmann (1932b) RO, CR.

Castilleja parviflora Bong. var. douglasii (Benth.) Jeps.
Castilleja affinis H. & A. ssp. affinis
Hoffmann (1932b) RO, CR.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 99

Castilleja sp.

Castilleja grisea Dunkle

Gentry (1949) based on Elmore 384 AHFH!, Elmore
411 LAM!, CL.

Castilleja wightii Elmer
Castilleja affinis H. & A. ssp. affinis
Smith (1976) RO?, CR, AN.

Castilleja wightii Elmer ssp. anacapensis (Dunkle) Penn.
Castilleja affinis H. & A. ssp. affinis
Abrams and Ferris (1923-1960) AN.

Catapodium rigidum (L.) C.E. Hubb.

Scleropoa rigida (L.) Griseb.

Thome (1967) CA.

Caulanthus lasiophyllus (H. & A.) Pays. var. inalienum Rob.
Caulanthus lasiophyllus (H. & A.) Pays.

Dunkle (1942) AN.

Caulanthus lasiophyllus (H. & A.) Pays. var. rigidum Rob.
Caulanthus lasiophyllus (H. & A.) Pays.

Dunkle (1942) AN.

Ceanothus arboreus Greene var. glabra Jeps.

Ceanothus arboreus Greene

Jepson (1925) RO; Munz (1935) RO; Eastwood (1941)
RO.

Ceanothus crassifolius Torr.

Ceanothus megacarpus Nutt. ssp. insularis (Eastw.) Raven
Greene (1887a) CR; Greene (1887b) MI; Brandegee
(1888) RO; Yates (1889) MI, RO, CR; Ford (1890)
CR; Brandegee (1890a) CA; Brandegee (1890b) MI,
RO, CR, CA; Jepson (1909-1943) CR; Millspaugh
and Nuttall ( 1 923) CA; Dunkle (1950) MI, RO, CR,
AN, CA.

Ceanothus crassifolius Torr. var. planus Abrams
incertae sedis
Jepson (1925) CR.

Ceanothus cuneatus (Hook.) Nutt.
incertae sedis

Watson ( 1876) GU; Eastwood (1929) GU; Dunkle (1950)
GU.

Ceanothus megacarpus Nutt. ssp. megacarpus
Trask (1899) CA in part.

Ceanothus insularis Eastw.

Ceanothus megacarpus Nutt. ssp. insularis (Eastw.) Raven
Abrams and Ferris ( 1 923-1960) RO, CR, CA; Eastwood
(1941) RO, CR, CA, CL; Munz and Keck (1959)
RO, CR, CA.

Ceanothus megacarpus Nutt. var. pendulus McMinn
Ceanothus megacarpus Nutt. ssp. megacarpus
McMinn (1942) based in part on Parish 10747 DS!, CA.
Ceanothus sorediatus H. & A.

Ceanothus arboreus Greene
Lyon (1886) CA.

Centaurium venustum (Gray) Rob.

Centaurium davyi (Jeps.) Abrams
Gentry (1949) based on Elmore 204 LAM!, misidenti-
fied, RO.

Centaurium davyi (Jeps.) Abrams
Clokey (193 1) based on Clokey 5117 NY!, misidentified,
CR; Eastwood (1941) CR; Gentry (1949) CR.

Centunculus minimus L.

Anagallis minima (L.) Krause

Hoffmann (1932b) RO; Munz (1935) RO; Eastwood
(1941) RO; Munz and Keck (1959) RO.

Cerastium viscosum L.

Cerastium glomeratum Thuill.

Hoffmann (1932a) MI, RO, CR; Munz (1935) MI, CL;
Eastwood (1941) MI, RO, CR, CA.

Cerasus ilicifolia Nutt, ex H. & A.

Prunus lyonii (Eastw.) Sarg.

Trask (1899) CA.

Cerasus ilicifolia Nutt, ex H. & A. var. integrifolia
Prunus lyonii (Eastw.) Sarg.

Trask (1899) CA.

Ceratochloa grandiflora Hook.

Bromus carinatus H. & A. (Raven, 1963).

Lyon (1886) CL.

Cercocarpus betulaefolius Nutt, ex Hook.

Cercocarpus betuloides Nutt, ex T. & G. ssp. blancheae
(C.K. Schneid.) Thome
Greene (1887a) CR; Yates (1889) CR.

Cercocarpus betuloides Nutt, ex T. & G. var. alnifolius (Rydb.)
Dunkle

Cercocarpus betuloides Nutt, ex T. & G. ssp. blancheae
(C.K. Schneid.) Thome
Dunkle (1950) RO, CR, CA.

Cercocarpus betuloides Nutt, ex T. & G. var. multiflorus Jeps.
Cercocarpus betuloides Nutt, ex T. & G. ssp. betuloides
Hoffmann (1932b) CR; Munz (1935) CR; Eastwood
(1941) CR; Dunkle (1950) CR.

Cercocarpus betuloides Nutt, ex T. & G. ssp. blancheae
(C.K. Schneid.) Thome

Jepson (1909-1943) CA; Jepson (1925) CA; Eastwood
(1941) CA; Dunkle (1950) CA.

Cercocarpus betuloides Nutt, ex T. & G. var. traskiae (Eastw.)
Dunkle

Cercocarpus traskiae Eastw.

Dunkle (1950) CA; Munz and Keck (1959) CA.
Cercocarpus parvifolius Nutt.

Cercocarpus betuloides Nutt, ex T. & G. ssp. blancheae
(C.K. Schneid.) Thome

Lyon (1886) CA; Brandegee (1890b) CR, CA; Trask
(1899) CA.

Cercocarpus traskiae Eastw.

Cercocarpus betuloides Nutt, ex T. & G. ssp. blancheae
(C.K. Schneid.) Thome

Munz (1935) probably based on Wolf 2752 RSA!, mis-
identified, CR.

Cereus sp.
incertae sedis
Yates (1889) AN.

Cereus emoryi Engelm.

Bergerocactus emoryi (Engelm.) Britt. & Rose

Lyon (1886) CL; Brandegee (1890a) CA; Brandegee
(1890b) CA, CL; Trask (1899) CA; Trask (1904)
CL; Jepson ( 1 909-1943) CA, CL; Jepson (1925) CL;
Munz (1935) CA, CL; Eastwood (1941) CA, CL;

100 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Dunkle (1950) CA, CL; Munz and Keck (1959) CA,
CL.

Chaenactis tenuifolia Nutt.

Chaenactis glabriscula DC. var. lanosa (DC.) Hall
Brandegee (1888) RO; Yates (1889) RO; Brandegee
(1890b) RO; Eastwood (1941) RO.

Chaetopappa lyonii (Gray) Keck
Pentachaeta lyonii Gray

Abrams and Ferris (1923-1960) CA; Munz and Keck
(1959) CA; Munz (1974) CA.

Cheilanthes californica (Hook.) Mett.
omitted as column transposition for CA.

Dunkle (1950, p. 293) BA.

Aspidotis californica (Hook.) Nutt, ex Copel.

Greene (1887a) CR; Yates (1889) CR; Yates (1890) CR;
Brandegee (1890a) CA; Brandegee (1890b) CA;
Trask (1899) CA; Millspaugh and Nuttall (1923)
CA; Clokey (1931) CR; Hoffmann (1932a) CR; Munz
(1935) CR, CA; Dunkle (1940a) CR, CA; Eastwood
(1941) CR, CA; Dunkle (1950) CR.

Cheilanthes clevelandii D.C. Eat.

Dunkle (1940a) RO, CR; Eastwood (1941) RO, CR;
Dunkle (1950) RO, CR.

Cheilanthes myriophylla Desv.

Cheilanthes clevelandii D.C. Eat.

Brandegee (1888) based on Brandegee s.n. in 1888 UC!,
misidentified, RO, based on Brandegee s.n. in Apr.
1 888 UC!, misidentified, CR; Yates ( 1 889) CR; Yates
(1890) CR; Brandegee (1890b) RO, CR.

Cheilanthes newberryi (D.C. Eat.) Domin
Notholaena newberryi D.C. Eat.

Munz and Keck (1959) CL; Raven (1963) CL.

Cheiranthus asper Cham. & Schlecht.
incertae sedis

Brandegee (1888) RO; Brandegee (1890b) RO.

Chenopodium album L.

Chenopodium murale L.

Watson (1876) based on Palmer s.n. in 1875 NY!, mis-
identified, GU; Lyon (1886) (possibly according to
Raven, 1963), CL; Dunkle (1942) probably based
on Dunkle 7610 LAM!, misidentified, AN; Yates
(1889) MI.

Chenopodium berlandieri Moq. ssp. zschakei (J. Murr.) Zo-
bel.

Chenopodium berlandieri Moq. var. sinuatum (J. Murr.)

H.A. Wahl
Thome (1967) CA.

Chorizanthe wheeled Wats,
omitted as unsubstantiated on CA and CL.

Eastwood (1941) CA, CL.

Cirsium coulteri Harv. & Gray in Gray
Circium proteanum J.T. Howell
Eastwood (1941) RO, CR.

Cirsium occidental (Nutt.) Jeps. var. coulteri (Harv. & Gray
in Gray) Jeps.

Cirsium proteanum J.T. Howell
Hoffmann (1932b) RO, CR.

Cirsium proteanum J.T. Howell
omitted as unsubstantiated on CR.

Smith (1976) CR “?.”

Cirsium undulatum (Nutt.) Spreng.

Cirsium ochrocentrum Gray
Thome (1967) based on Eastwood 6510 CAS!, US!, mis-
identified, CA.

Claytonia perfoliata Donn
incertae sedis

Greene (1887a) CR; Brandegee (1888) RO; Yates ( 1 889)
RO, CR.

Clematis ligusticifolia Nutt, in T. & G.
omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Eastwood (1941) CL.

Clematis pauci flora Nutt, in T. & G.
omitted as unsubstantiated on CA and CL; also omitted
by Raven (1963).

Eastwood (1941) CA, CL.
probably Clematis lasiantha Nutt, in T. & G.

Brandegee (1890b) CR; Eastwood (1941) RO; Smith
(1976) based on Howell 6217 CAS!, misidentified,
CR.

Cneoridium dumosum (Nutt.) Hook.

Lycium californicum Nutt.

Munz (1974) in part, misidentified, CL.

Cnicus sp.

possibly Cirsium occidentale (Nutt.) Jeps.

Greene (1887b) MI.

Cnicus lilacinus Greene

Cirsium occidentale (Nutt.) Jeps.

Greene (1887a) CR; Yates (1889) CR; Eastwood (1941)
RO, CR.

Cnicus occidentalis (Nutt.) Gray
Cirsium occidentale (Nutt.) Jeps.

Lyon (1886) CA; Greene (1887b) MI; Brandegee (1888)
RO; Yates (1889) MI; Brandegee (1890b) MI, RO,
CR, CA; Trask (1904) CL.

Collomia gilioides Benth. var. glutinosa Gray
Allophyllum gilioides (Benth.) A. Grant & V. Grant
Watson (1876) based on Palmer 77 MO!, NY!, mis-
identified, GU; Eastwood (1929) GU.
Comarostaphylis polifolia (HBK.) Zucc.

Comarostaphylis diversifolia (Parry) Greene ssp. planifolia
(Jeps.) Wallace ex Thome

Millspaugh and Nuttall (1923) CA; Eastwood (1941) CA.
Convolvulus aridus Greene ssp. intermedius Abrams

Calystegia macrostegia (Greene) Brummitt ssp. intermedia
(Abrams) Brummitt
Munz and Keck (1959) CA.

Convolvulus californicus Choisy
Calystegia macrostegia (Greene) Brummitt ssp. macro-
stegia

Lyon (1886) CA; Brandegee (1890b) CA; Davidson
( 1 896) CA; Millspaugh and Nuttall ( 1 923) CA; East-
wood (1941) CA.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 101

Convolvulus cyclostegius House

Calystegia macrostegia (Greene) Brum mitt ssp. cyclostegia
(House) Brummitt

Eastwood (1941) CA; Munz and Keck (1959) CA.

Convolvulus macrostegius Greene

Calystegia macrostegia (Greene) Brummitt ssp. amplissi-
ma Brummitt

Lyon ( 1 886) CL; Eastwood (1898) NI; Trask ( 1 904) CL;
Jepson (1909-1943) BA, CL; Abrams and Ferris
(1923-1960) NI, BA, CL; Jepson (1925) CL; East-
wood (1941) NI, BA, CL; Munz and Keck (1959)
NI, BA, CL; Raven (1963) CL.

Calystegia macrostegia (Greene) Brummitt ssp. macro-
stegia

Greene (1885) GU; Lyon (1886) CA; Greene (1887a)
CR; Greene (1887b) MI; Brandegee (1888) RO;
Yates (1889) MI, RO, CR, AN; Franceschi (1893)
GU; Davidson (1896) CA; Jepson (1909-1943) CR,
AN, GU; Abrams and Ferris (1923-1960) CR, AN,
GU; Jepson (1925) CR, GU; Eastwood (1929) GU;
Eastwood (1941) MI, RO, CR, AN; Howell (1942)
GU; Munz and Keck (1959) RO, CR, CA, GU.

Convolvulus occidentalis Gray

Calystegia macrostegia (Greene) Brummitt ssp. amplissi-
ma Brummitt

Brandegee (1890b) CL.

Calystegia macrostegia (Greene) Brummitt ssp. macro-
stegia

Watson (1876) GU; Lyon ( 1 886) CA; Brandegee ( 1 890b)
MI, RO, CR, CA; Trask ( 1 889) CA; Millspaugh and
Nuttall (1923) CA.

Convolvulus occidentalis Gray var. cyclostegius (House) Jeps.

Calystegia macrostegia (Greene) Brummitt ssp. amplissi-
ma Brummitt

Gentry (1949) based on Elmore 313 AHFH!, misiden-
tified, BA, based on Elmore 396 AHFH!, misiden-
tified, CL.

Calystegia macrostegia (Greene) Brummitt ssp. cyclostegia
(House) Brummitt

Munz ( 1 935) CA; Gentry ( 1 949) CA; Dunkle (1950) CA.

Calystegia macrostegia (Greene) Brummitt ssp. macro-
stegia

Gentry (1949) based on Elmore 282 AHFH!, misiden-
tified, CR.

Convolvulus occidentalis Gray var. macrostegius Munz

Calystegia macrostegia (Greene) Brummitt ssp. amplissi-
ma Brummitt

Munz (1935) CL; Dunkle (1942) BA; Dunkle (1950) NI,
BA, CL.

Calystegia macrostegia (Greene) Brummitt ssp. macro-
stegia

Munz (1935) CR, CA, GU; Dunkle (1942) AN; Dunkle
(1950) MI, RO, CR, AN, CA, GU.

Convolvulus sepium L.

Calystegia macrostegia (Greene) Brummitt ssp. intermedia
(Abrams) Brummitt

Millspaugh and Nuttall (1923) based in part on Mill-
spaugh 4657 F!, misidentified, CA; Eastwood ( 1 94 1 )
CA.

Convolvulus soldanella L.

Calystegia soldanella (L.) R. Br.

Lyon ( 1 886) CA; Brandegee ( 1 890a) CA; Millspaugh and
Nuttall (1923) CA; Hoffmann (1932b) MI, CR;
Eastwood (1941) MI, CR, CA; Raven (1963) CL.

Coreopsis gigantea (Kell.) Hall
omitted as unsubstantiated~on CL; also omitted by Raven
(1963).

Munz (1935) “all our islands.”

Corethrogyne filaginifolia (H. & A.) Nutt. var. latifolia Hall
Corethrogyne filaginifolia (H. & A.) Nutt. ssp. filaginifolia
Dunkle (1942) AN; Dunkle (1950) AN; Munz and Keck
(1959) AN; Munz (1974) AN.

Corethrogyne filaginifolia (H. & A.) Nutt. var. robusta Greene
Corethrogyne filaginifolia (H. & A.) Nutt. ssp. filaginifolia
Abrams and Ferris (1923-1960) MI, RO, CR; Eastwood
(1941) MI, RO, CR; Dunkle (1942) AN; Gentry
(1949) based on Elmore 220 AHFH!, misidentified,
AN, and MI, RO, CR; Dunkle (1950) MI, RO, CR,
AN; Munz and Keck (1959) MI, RO; Munz (1974)
MI, RO.

Corethrogyne filaginifolia (H. & A.) Nutt. var. virgata (Benth.)
Gray

Corethrogyne filaginifolia (H. & A.) Nutt. ssp. filaginifolia
Eastwood (1941) MI, RO, CR; Dunkle (1950) MI, RO,
CR, CA; Munz and Keck (1959) CA.

Corethrogyne lavendulacea Greene
Corethrogyne filaginifolia (H. & A.) Nutt. ssp. filaginifolia
Eastwood (1941) CA.

Cornus glabrata Benth. var. catalinensis (Millsp.) Dunkle
Cornus glabrata Benth.

Dunkle (1950) CA.

Cortaderia selloana (Schult.) Asch. & Graebn.

Cortaderia atacamensis (Phil.) Pilg.

Thome (1967) CA.

Cotyledon sp.
incertae sedis
Franceschi (1893) GU.

Dudleya virens (Rose) Moran
Lyon (1886) CL.

Cotyledon caespitosa Haw.

Dudleya greenei Rose

Lyon (1886) CA; Brandegee (1890b) CR, CA; Davidson
(1896) CA; Trask (1899) CA; Millspaugh and Nut-
tall (1923) CA.

Cotyledon lanceolata (Nutt, ex T. & G.) Benth. & Hook. ex.
Wats.

Dudleya greenei Rose

Greene (1887a) CR; Greene (1887b) MI; Brandegee
(1888) RO; Yates (1889) MI, RO, CR, AN; Bran-
degee (1890b) MI, RO, CR.

Dudleya virens (Rose) Moran

Brandegee (1890a) CA; Brandegee (1890b) CA; Trask
(1899) CA.

102 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Cotyledon laxa (Lindl.) Benth. & Hook, ex Wats.
incertae sedis

Greene ( 1 887a) CR; Yates ( 1 889) CR; Brandegee ( 1 890b)
CR.

Cressa cretica L.

Cressa truxillensis HBK. var. vallicola (Heller) Munz
Lyon (1886) CA; Greene (1887b) MI; Yates (1889) MI;
Brandegee (1890b) MI, CA; Jepson (1909-1923) MI,
CA; Hoffmann (1932b) RO, CR.

Crossostephium insulare Rydb.

Artemisia nesiotica Raven

Davidson and Moxley (1923) NI, CL.

Crvptantha ambigua (Gray) Greene
incertae sedis

Eastwood (1941) MI, CR, CA.

Cryptantha clevelandii Greene var. hispidissima Jtn.
Crvptantha clevelandii Greene var. clevelandii
Dunkle (1942) AN, BA; Dunkle (1950) MI, RO, CR,
BA, CL, GU.

Cryptantha intermedia (Gray) Greene
omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Dunkle (1942) BA.

Cryptantha clevelandii Greene var. florosa Jtn.

Foreman (1967) based on Kanakoff s.n. Apr. 12, 1940
LAM!, misidentified, NI.

Cryptantha leiocarpa (F. & M.) Greene
Cryptantha clevelandii Greene var. florosa Jtn.
Millspaugh and Nuttall (1923) based on Hasse 4156
NY!, misidentified, CA.

Cryptantha ramosissima (Gray) Greene
Cryptantha maritima (Greene) Greene

Millspaugh and Nuttall (1923) CA; Eastwood (1941) CA.
Cryptantha torreyana (Gray) Greene
Cryptantha maritima (Greene) Greene
Eastwood (1898) based on Trask 57 LAM!, GH!, NY!,
misidentified, NI.

Cryptantha traskiae Jtn.
omitted as unsubstantiated on CA.

Eastwood (1898) CA.
omitted as column transposition for CA.

Dunkle (1950, p. 293) BA.

Cryptantha clevelandii Greene var. clevelandii
Dunkle (1942) probably based on Dunkle 7446b AHFH!,
misidentified, BA; also noted by Philbrick (1972).
Cupressus macrocarpa Hartw.

Cupressus guadalupensis Wats. ssp. gnadalupensis
Watson (1876) GU.

Cuscuta californica H. & A.
omitted as unsubstantiated on CR.

Smith (1976) CR.

Cuscuta subinclusa Dur. & Hilg.

Cuscuta ceanothi Behr

Greene (1887a) CR; Yates (1889) CR; Brandegee (1890b)
CR; Jepson (1909-1943) CR; Eastwood (1941) CR.
Cytisus canariensis Steud.

Cytisus monspessulans L.

Millspaugh and Nuttall (1923) CA; Eastwood (1941 )CA.

Datura meteloides A. DC.

Datura wrightii Regel

Greene (1887a) CR; Yates (1889) CR; Brandegee ( 1 890a)
CA; Brandegee ( 1 890b) CR, CA; Jepson ( 1 909-1943)
CR; Millspaugh and Nuttall (1923) CA; Hoffmann
(1932b) RO; Eastwood (1941) RO, CR, CA; Dunkle
(1950) RO, CR, CA; Smith (1976) RO, CR.
Deinandra paniculata (Gray) Davids. & Mox.

Hemizonia increscens (Hall ex Keck) Tanowitz ssp. in-
crescens

Davidson and Moxley (1923) RO.

Deinandra wrightii (Gray) Greene
probably Hemizonia fasciculat a (DC.) T. & G.

Abrams (1917) CA.

Delphinium sp.

probably Delphinium parryi Gray ssp. parryi
Greene (1887a) CR; Brandegee (1890a) CA.
Delphinium hesperium Gray
Delphinium parryi Gray

Millspaugh and Nuttall (1923) CA; Eastwood (1941) CA.
Dendromecon rigidus Benth. var. harfordii K. Bdg.

Dendromecon rigidus Benth. ssp. rhamnoides (Greene)
Thome

Jepson (1909-1943) probably misidentified, CA.
Dendromecon arborea Greene

Dendromecon rigidus Benth. ssp. rhamnoides (Greene)
Thome

Millspaugh and Nuttall (1923) CA; Eastwood (1941 )CA.

Dendromecon densifolia Greene

Dendromecon rigidus Benth. ssp. harfordii (Kell.) Raven
Eastwood (1941) RO.

Dendromecon flexile Greene

Dendromecon rigidus Benth. ssp. harfordii (Kell.) Raven
Greene (1887a) CR; Yates (1889) CR; Ford (1890) CR;
Eastwood (1941) CR.

Dendromecon rigidus Benth. ssp. rhamnoides (Greene)
Thome

Davidson ( 1 896) CA; Millspaugh and Nuttall ( 1 923) CA.
Dendromecon harfordii Kell.

Dendromecon rigidus Benth. ssp. harfordii (Kell.) Raven
Kellogg (1873) RO; Ford ( 1 890) CR; Munz (1935) Santa
Barbara Isl.; Eastwood (1941) RO; Dunkle (1950)
RO, CR; Munz and Keck (1959) RO, CR.
Dendromecon rigidus Benth. ssp. rhamnoides (Greene)
Thome

Trask (1899) CA; Davidson and Moxley (1923) CA;
Eastwood (1941) CA; Dunkle (1950) CA, CL.
Dendromecon harfordii Kell. var. rhamnoides (Greene) Munz
Dendromecon rigidus Benth. ssp. rhamnoides (Greene)
Thome

Munz and Keck (1959) CA, CL.

Dendromecon rhamnoides Greene
Dendromecon rigidus Benth. ssp. rhamnoides (Greene)
Thome

Millspaugh and Nuttall ( 1 923) CA; Eastwood (1941) CA,
CL.

Dendromecon rigidus Benth. ssp. rigidus

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 103

Dendromecon rigidus Benth. ssp. rhanmoides (Greene)
Thome

Raven (1963) CL.

Dentaria californica Nutt.

Cardamine californica (Nutt.) Greene
Eastwood (1941) MI, RO, CR, CA.

Dentaria integrifolia Nutt. var. californica Jeps.

Cardamine californica (Nutt.) Greene
Jepson (1909-1943) CR; Hoffmann (1932b) MI; Smith
(1976) all four Channel Islands.

Descurainia pinnata (Walt.) Britt, ssp. halictorum (Ckll.) Detl.
Descurainia pinnata (Walt.) Britt, ssp. menziesii (DC.) Detl.
Wiggins (1980) probably misidentified, GU.
Dichelostemma capitatum (Benth.) Wood
Dichelostemma pulchellum (Salisb.) Heller
Millspaugh and Nuttall (1923) CA.

Dichondra argentea Willd.

Dichondra occidentalis House

Brandegee (1888) RO; Yates (1889) RO.

Dichondra donelliana Tharp & Johnst.

Dichondra occidentalis House
Tharp and Johnston (1961) based in part on Wolf 2841
POM!, misidentified, CR; Smith (1976) MI, RO,
CR.

Dichondra repens Forst.

Dichondra occidentalis House

Brandegee ( 1 890a) CA; Brandegee ( 1 890b) RO, CA; Da-
vidson (1896) CA; Jepson (1909-1943) RO, CA;
Hoffmann (1932b) CR; Eastwood (1941) RO, CR,
CA.

Diplacus arachnoideus Greene
Diplacus longiflorus Nutt. ssp. longiflorus
Greene (1887a) CR; Yates (1889) CR; Eastwood (1941)
RO, CR; McMinn (1951) CR.

Diplacus linearis (Benth.) Greene
Diplacus longiflorus Nutt. ssp. longiflorus
Eastwood (1941) RO.

Diplacus puniceus Nutt.

Millspaugh and Nuttall ( 1923)CA; Eastwood ( 1941)CA.
Diplacus longiflorus Nutt.

omitted as unsubstantiated on CA.

Gentry (1949) CA.

Diplacus parviflorus Greene
Gentry (1949) probably based on Elmore 185 AHFH!,
misidentified, RO.

Diplostephium canum Gray
Haplopappus canus (Gray) Blake
Watson (1876) GU; Brandegee (1890b) GU; Franceschi
(1893) GU.

Haplopappus detonus (Greene) Raven

Brandegee (1888) RO; Brandegee (1890b) RO, CR.
Dissanthelium californicum (Nutt.) Benth.
omitted as unsubstantiated on RO and CR.

Eastwood (1941) RO, CR.

Distichlis dentata Rydb.

Distichlis spicata (L.) Greene var. stolonifera Beetle
Cockerell (1937) MI.

Distichlis maritima Raf.

Distichlis spicata (L.) Greene var. stolonifera Beetle
Brandegee ( 1 890a) CA; Brandegee ( 1 890b) MI, RO, CR,
CA.

Distichlis stricta (Torr.) Rydb. var. laxa (Vasey) Fawcett &
West ex Munz

Distichlis spicata (L.) Greene var. stolonifera Beetle
Munz (1 935) RO, CR, CA, CL; Dunkle ( 1 942) AN; Dun-
kle (1950) MI, RO, CR, AN, NI, CA, CL.
Dithyraea californica Harv. var. maritima (A. Davids.) A.
Davids, ex Rob. in Gray
Dithyraea maritima A. Davids.

Eastwood (1898) NI; Hoffmann (1932b) MI; Eastwood
(1941) MI, NI.

Dithyraea maritima A. Davids,
omitted as unsubstantiated on CA.

Davidson and Moxley (1923) CA.

Dodecatheon clevelandii Greene ssp. sanctarum (Greene)
Abrams

Dodecatheon clevelandii Greene ssp. insularis H.J. Thomps.
Abrams and Ferris (1923-1960) CR, AN, CA, CL.
Dodecatheon hendersoni Gray
Dodecatheon clevelandii Greene ssp. insularis H.J. Thomps.
Davidson (1896) CA; Brandegee (1888); RO; Yates
(1889) RO; Millspaugh and Nuttall (1 923) CA; Munz
(1935) RO, CA; Eastwood ( 1 94 1 ) CA; Dunkle ( 1 942)
AN.

Dodecatheon jejfreyi Moore

Dodecatheon clevelandii Greene ssp. insularis H.J. Thomps.
Davidson (1896) CA; Greene (1887a) CR; Yates (1889)
CR.

Dodecatheon meadia L.

Dodecatheon clevelandii Greene ssp. insularis H.J. Thomps.
Watson (1876) GU; Greene (1885) GU; Brandegee
( 1 890a) CA; Brandegee ( 1 890b) RO, CR, CA; Fran-
ceschi (1893) GU.

Dudleya farinosa (Lindl.) Britt. & Rose
Dudleya greenei Rose

Gentry (1949) based on Elmore 314 LAM!, misidenti-
fied, MI.

Dudleya hassei (Rose) Moran

omitted as unsubstantiated on GU.

Moran (1959) GU.

Dudleya virens (Rose) Moran
Dudleya hassei (Rose) Moran

Munz and Keck (1959) CA, in part.

Echeveria albida (Rose) Berger in Engl. & Prantl
Dudleya greenei Rose

Munz (1935) MI; Dunkle (1950) MI.

Dudleya traskiae (Rose) Moran

Munz (1935) BA; Dunkle ( 1 942) BA; Dunkle (1950) BA.
Dudleya virens (Rose) Moran

Munz (1935) CA in part, CL; Dunkle (1950) CA in part,
CL, GU.

Echeveria caespitosa (Haw.) DC.

Dudleya caespitosum (Haw.) Britt.

Jepson (1909-1943) CR.

104 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Echeveria greenei (Rose) Berger in Engl. & Prantl

omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Dunkle (1942) BA.
omitted as unsubstantiated on GU.

Dunkle (1950) GU.

Echeveria lanceolata Nutt, ex T. & G.

Dudley a virens (Rose) Moran
Jepson (1909-1943) CA.

Echeveria viscida (Wats.) Berger in Engl. & Prantl var. in-
sulare (Rose) Jeps.

Dudleya hassei (Rose) Moran
Jepson (1909-1943) CA in part.

Dudleya traskiae (Rose) Moran
Jepson (1909-1943) BA.

Dudleya virens (Rose) Moran
Jepson (1909-1943) CA in part, CL.

Echidocarya californica Gray
Plagiobothrys californicus (Gray) Greene var. gracilis Jtn.
Jepson (1909-1943) CR, CA, CL.

Echidocarya californica Gray ssp. fulvescens (Jtn.) Abrams
Plagiobothrys californicus (Gray) Greene var. fulvescens
Jtn.

Abrams and Ferris (1923-1960) RO, AN, CA.
Echinocystis fabacea Naud.

Marah macrocarpus (Greene) Greene

Brandegee (1888) RO; Brandegee (1890b) MI, RO, CR,
CA, CL; Hoffmann ( 1 932b) MI, RO, CR, AN; Munz
(1935) MI; Dunkle (1942) AN; Dunkle (1950) MI.
Echinocystis guadalupensis (Wats.) Naud.

Marah guadalupensis (Wats.) Greene
Greene ( 1 8 8 5) GU; Franceschi ( 1 8 9 3) GU; Dunkle (1950)
GU.

Marah macrocarpus (Greene) Greene
Greene (1887a) CR; Greene (1887b) MI; Yates (1889)
MI, CR; Dunkle (1950) MI, RO, CR, AN.
Eleocharis mamillata Lindb. f.

E/eocharis macrostachya Britt, in Small
Munz (1935) RO, CA.

Eleocharis palustris (L.) R. & S.

Eleocharis macrostachya Britt, in Small

Brandegee (1890a) CA; Brandegee (1890b) CA; David-
son (1896) CA; Eastwood (1898) NI, CR; Mills-
paugh and Nuttall (1923) CA; Hoffmann (1932a)
RO; Eastwood (1941) RO, NI, CA; Raven (1963)
CL; Thome (1967) CA.

Elymus multisetus (J.G. Sm.) Jones
Sitanion jubatum J.G. Sm.

Thome (1967) based on Nuttall 314 F!, misidentified,
CA.

Elymus triticoides Buckl. ssp. multiflorus Gould
Elymus triticoides Buckl.

Smith (1976) islands.

Epilobium adenocaulon Hausskn.

Epilobium ciliatum Raf. ssp. ciliatum

McClatchie (1894) CA; Eastwood (1941) CR; Smith
(1976) MI, CR.

Epilobium adenocaulon Hausskn. var. holosericeum (Trel.)
Munz

Epilobium ciliatum Raf. ssp. ciliatum
Thorne (1967) CA.

Epilobium adenocaulon Hausskn. var. parishii (Trel.) Munz
Epilobium ciliatum Raf. ssp. ciliatum
Smith (1976) CR.

Epilobium californicum Hausskn.

Epilobium ciliatum Raf. ssp. ciliatum
Clokey (1931) CR.

Epilobium coloratura Muhl.

Epilobium ciliatum Raf. ssp. ciliatum
Greene (1887a) CR; Yates (1889) CR; Brandegee (1890b)
CR; Eastwood (1941) CR, CA.

Epilobium holosericeum Trel.

Epilobium ciliatum Raf. ssp. ciliatum

Millspaugh and Nuttall (1923) CA; Eastwood (1941) CA.
Epilobium minutum Lindl. ex Hook.

Epilobium foliosum (T. & G.) Suksd. (Seavey et af, 1 977).
Watson (1876) based on Palmer 31 MO! (collection
number reported by Seavey, Wright and Raven, 1977
is in error), misidentified, GU; Greene (1885) GU;
Eastw'ood (1929) GU.

Equisetum sp.
incertae sedis
Greene (1887a) CR.

Equisetum funstoni A. A. Eat.

Equisetum laevigatum A. Br.

Hoffmann (1932a) CR; Eastwood (1941) CR, CA; Munz
and Keck (1959) CA.

Equisetum hyemale L. var. californicum Milde.

Equisetum hyemale L. var. affine (Engelm.) A. A. Eat.
Hoffmann (1932a) CR; Munz ( 1 93 5) CR; Dunkle (1950)
CR; Munz and Keck (1959) CR.

Equisetum kansanum J.H. Schaffn.

Equisetum laevigatum A. Br.

Hoffmann (1932a) CR; Millspaugh and Nuttall (1923)
CA; Eastwood (1941) CR, CA.

Equisetum mexicanum Milde.

probably Equisetum laevigatum A. Br.

McClatchie (1894) CA.

Equisetum robustum A. Br.

Equiseteum laevigatum A. Br.

Davidson (1894) CA.

Erigeron foliosus Nutt.
incertae sedis

Lyon (1886) CA; Brandegee (1888) RO; Brandegee
(1890b) MI, RO, CR, CA.

Eriodictyon tomentosum Benth.

Eriodictyon traskiae Eastw. ssp. traskiae

Lyon (1886) CA; Brandegee (1890b) CA; Abrams and
Ferris (1923-1960) CA.

Eriogonum arborescens Greene
omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Davidson and Moxley (1923) BA.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 105

Eriogonum giganteum Wats.

Eriogonum giganteum Wats. ssp. formosum (K. Bdg.) Ra-
ven

Jepson (1925) CL; Munz (1935) CL; Gentry (1949) CL.
introduced on CR.

Jepson (1925) CR; Gentry (1949) CR.

Eriogonum giganteum Wats. var. compactum Dunkle
omitted as unsubstantiated on CA.

Dunkle (1950) CA.

Eriogonum giganteum Wats. var. formosum K. Bdg.
omitted as unsubstantiated on RO.

Eastwood (1941) RO.

Eriogonum grande Greene
omitted as column transposition for CA.

Dunkle (1950) BA.

omitted as unsubstantiated on MI and BA.

Gentry (1949) MI, BA.

Eriogonum latifolium Sm.

Eriogonum grande Greene ssp. grande
Yates (1889) AN.

Eriogonum latifolium Sm. ssp. grande (Greene) S. Stokes
Eriogonum grande Greene ssp. grande
Davidson and Moxley (1923) CA; Munz and Keck (1959)
CR, AN, CA, CL; Raven ( 1 963) CL; Foreman (1967)
NI; Thome (1967) CA.

Eriogonum latifolium Sm. var. rubescens (Greene) Munz
Eriogonum grande Greene ssp. rubescens (Greene) Munz
Munz and Keck (1959) MI, RO, CR.

Eriogonum molle Greene

Eriogonum zapatoense Moran
Moran (1951) GU.

Eriogonum nudum (Dougl. ex Benth.) S. Stokes
Eriogonum grande Greene ssp. grande

Trask (1899) CA; Davidson (1896) CA; Brandegee
(1890b) CA, CL; Trask (1904) CL; Abrams (1917)
CA; Millspaugh and Nuttall (1923) CA.
Eriogonum grande Greene ssp. rubescens (Greene) Munz
Brandegee (1888) RO; Brandegee (1890b) MI, RO, CR.
Eriogonum nudum (Dougl. ex Benth.) S. Stokes var. grande
Jeps.

Eriogonum grande Greene ssp. grande
Jepson (1909-1943) CR; Hoffmann (1932a) RO in part,
CR in part, AN; Dunkle (1942) AN.

Eriogonum grande Greene ssp. rubescens (Greene) Munz
Jepson (1909-1943) MI, CR in part; Hoffmann (1932a)
MI, RO in part, CR in part.

Eriogonum nudum (Dougl. ex Benth.) S. Stokes var. pauci-
florum Wats.

Eriogonum grande Greene ssp. grande
Lyon (1886) CA, CL.

Eriogonum sp.

Eriogonum grande Greene ssp. formosum (K. Bdg.) Raven
Lyon (1886) CL.

Eriogonum rubescens Greene

omitted as unsubstantiated on CA and CL.

Eastwood (1941) CA, CL; Gentry (1949) CA, CL.

Eriogonum grande Greene ssp. rubescens (Greene) Munz
Greene (1887b) MI; Yates (1889) MI, RO, CR; East-
wood (1941) MI, RO, CR; Gentry (1949) MI, RO,
CR.

Eriophyllum sp.

Eriophyllum lanatum (Pursh) Forbes var. grandiflorum
(Gray) Jeps.

Franceschi (1893) GU.

Eriophyllum staechadifolium Lag.
incertae sedis

Brandegee (1888) RO; Yates (1889) RO, CR; Brandegee
(1890b) RO, CR.

Eriophyllum staechadifloium Lag. var. artemisiaefolium
(Less.) Macbr.

omitted as unsubstantiated on CA.

Dunkle (1950) CA.

Eritrichum angustifolium Torr.

Cryptantha maritima (Greene) Greene
Watson (1876) based on Palmer 67 MO!, GU.
Erodium botrys (Cav.) Bertol.

omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Dunkle (1942) BA.

Erodium brachycarpum (Gordon) Thell.

Erodium obtusiplicatum (Maire, Weiller & Wilcz.) J.T.
Howell

Smith (1976) RO.

Erysimum asperum DC.
incertae sedis

Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890b) CR; Jepson (1909-1943) CR; Eastwood
(1941) RO, CR, CA.

Erysimum insulare Greene

Hoffmann (1932b) based on Hoffmann s.n. SBM#141!,
RO, based on SBM#4151!, AN; Eastwood (1941)
AN; Dunkle (1942) AN.

Erysimum capitatum (Dougl.) Greene
Erysimum ammophilum Heller

Munz (1935) probably based on Munz & Crow 11757
LA!, misidentified, RO.

Erysimum insulare Greene

omitted as unsubstantiated on NI and CA; also omitted
by Foreman (1967).

Eastwood (1941) NI, CA.

Erysimum morani Roll.

Moran (1951) GU.

Eschscholzia sp.

Eschscholzia californica Cham. var. californica

Hoffmann (1932b) based on Hoffmann s.n. SBM# 12039!,
CR.

Eschscholzia californica Cham.
incertae sedis

Brandegee (1888) RO; Brandegee (1890b) MI, RO, CR,
CA; Franceschi (1893) GU; Trask (1899) CA.
Eschscholzia californica Cham. var. californica
Greene (1885) GU; Gentry (1949) based on Elmore 187
AHFH! RO.

106 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Eschscholzia ramosa Greene
Brandegee (1890b) CL.

Eschscholzia californica Cham. var. hypecoides Gray
Eschscholzia elegans Greene
Watson (1876) based on Palmer 3 CM!, NY! in part,
US!, GU.

Eschscholzia ramosa Greene

Watson (1876) based on Palmer 3 NY! in part, GU.
Eschscholzia crossophylla Greene
Eschscholzia ramosa Greene
Eastwood (1941) CA.

Eschscholzia elegans Greene
Eschscholzia ramosa Greene
Jepson (1909-1943) CR, BA, CA; Millspaugh and Nut-
tall (1923) CA; Abrams and Ferris (1932-1960) is-
lands from RO to CL; HofFmann (1932b) RO, CR;
Munz (1935) CR, CA, CL; Eastwood (1941) MI,
RO, CR, CA, CL; Dunkle (1942) AN; Dunkle (1950)
MI, RO, CR, AN, NI, BA, CA, CL; Munz and Keck
(1959) Channel Islands.

Eschscholzia elegans Greene var. ramosa Greene
Eschscholzia ramosa Greene

Lyon (1886) CL; Greene (1885) GU; Franceschi (1893)
GU.

Eschscholzia glauca Greene
Eschscholzia californica Cham. var. californica
Greene (1887a) CR; Yates (1889) RO, CR; HofFmann
(1932b) CR; Eastwood (1941) CR.

Eschscholzia robusta Greene
incertae sedis
Eastwood (1941) RO.

Eschscholzia trichophylla Greene
Eschscholzia ramosa Greene
Eastwood (1941) CR.

Eschscholzia wrigleyana Millsp. & Nutt.

Eschscholzia californica Cham. var. peninsularis (Greene)
Munz

Millspaugh and Nuttall ( 1 923) CA; Eastwood (1941) CA.
Eulobus californicus Nutt, ex T. & G.

Camissonia californica (Nutt, ex T. & G.) Raven
Greene (1887a) CR; Yates (1889) CR; Brandegee ( 1 890a)
CA; Brandegee (1890b) CR, CA; Trask (1899) CA;
Millspaugh and Nuttall (1923) CA; Eastwood (1941)
RO, CR, CA.

Eunanus latifolius (Gray) Greene
Mimulus latifolius Gray
Greene (1885) GU.

Euphorbia misera Benth.

omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Dunkle (1950) BA.

Euryptera insularis (Eastw.) Coult. & Rose
Lomatium insulare (Eastw.) Munz
Davidson and Moxley (1923) NI.

Evax caulescens (Benth.) Gray var. humilus Jeps.

Evax sparsiflora (Gray) Jeps.

HofFmann (1932b) RO; Eastwood (1941) RO, CR.

Festuca bromoides L.

Vulpia bromoides (L.) S.F. Gray
Clokey (1931) CR; HofFmann ( 1 932a) MI, RO, CR; East-
wood ( 1 94 1 ) MI, RO, CR, NI, CA, CL; Raven (1963)
CL.

Festuca dertonensis (All.) Asch. & Graebn.

Vulpia bromoides (L.) S.F. Gray
Dunkle (1942) AN; Smith (1976) all four islands.

Festuca megalura Nutt.

Vulpia myuros (L.) K.C. Gmelin var. hirsuta Hack
Millspaugh and Nuttall (1923) CA; Clokey (1931) CR;
HofFmann (1932a) MI, RO, CR; Munz (1935) on
the islands; Eastwood (1941) MI, RO, CR, CA;
Dunkle ( 1 942) AN, BA; Howell ( 1 942) GU; Gentry
( 1 949) MI, RO, CR, CA; Raven (1963) CL; Thome
(1967) CA; Smith (1976) all four islands.

Festuca microstachys Nutt.

Vulpia microstachys (Nutt.) Benth. var. pauciflora (Beal)
Lonard & Gould

Watson (1876) GU; Greene (1885) GU; Brandegee (1888)
RO, CR; Yates (1889) RO, CR; Brandegee ( 1 890b)
RO, CR; Eastwood (1941) RO, CR.

Festuca myuros L.

incertae sedis

Greene (1887a) CR; Greene (1887b) MI; Yates (1889)
MI, CR; Brandegee (1890b) MI, CR.

Vulpia myuros (L.) K.C. Gmelin var. hirsuta Hack
Brandegee (1888) based on Brandegee s.n. in 1888
UC#121663!, RO; Yates (1889) RO; Brandegee
( 1 890b) RO; Vasey and Rose ( 1 890) GU; Eastwood
(1941) RO.

Vulpia myuros (L.) K.C. Gmelin var. myuros
Brandegee (1890a) CA; Jepson (1909-1943) CA; Mill-
spaugh and Nuttall (1923) CA; Abrams and Ferris
(1923-1960) CA; Eastwood (1941) CA; Thome
(1967) CA; Thome (1969) CL.

Festuca octoflora Walt.

Vulpia octoflora (Walt.) Rydb. var. hirtella (Piper) Henr.
Millspaugh and Nuttall (1923) CA; HofFmann (1932a)
MI, RO; Munz (1935) MI, RO; Dunkle (1942) AN;
Foreman (1967) NI; Thome (1967) CA; Smith
(1976) all four islands.

Festuca octoflora Walt. ssp. hirtella

Vulpia octoflora (Walt.) Rydb. var. hirtella (Piper) Henr.
Howell (1935) NI; Eastwood (1941) MI, RO, CR, NI,
CA; Dunkle (1942) AN; Dunkle (1950) MI, RO,
CR, AN, NI, CA; Raven (1963) CL.

Festuca pacifica Piper

Vulpia microstachys (Nutt.) Benth. var. pauciflora (Beal)
Lonard & Gould

HofFmann (1932a) MI, CR; Eastwood (1941) MI, CR,
NI; Gentry (1949) MI, CR, NI; Raven (1963) CL;
Smith (1976) AN.

Festuca pratensis Huds.

Festuca arundinacea Schreb.

Thome (1967) CA.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 107

Festuca reflexa Buckl.

Vulpia microstachys (Nutt.) Benth. var. pauciflora (Beal)
Lonard & Gould

Millspaugh and Nuttall (1923) CA; Eastwood (1941) CA;
Raven (1963) CL; Thome (1967) CA.

Festuca tenella Willd.

Vulpia octoflora (Walt.) Rydb. var. hirtella (Piper) Henr.
Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890a) based in part on Brandegee 49 US!, CA;
Brandegee ( 1 890b) CR, CA; Vasey and Rose ( 1 890)
based in part on Palmer 674 NY!, GU.

Filago arizonica Gray

Filago californica Nutt.

Vasey and Rose (1890) based on Palmer 895 ND-G!,
misidentified, GU; Eastwood (1941) possibly based
on Fosberg 7642 LAM!, misidentified, CA; Bunkle
(1950) CA.

Foeniculum foeniculum (L.) Karst.

Foeniculum vulgare Mill.

Millspaugh and Nuttall (1923) CA.

Foeniculum officinale All.

Foeniculum vulgare Mill.

Greene (1887a) CR; Yates (1889) CR; Brandegee ( 1 890b)
CR.

Frankenia grandiflora Cham. & Schlecht.

Frankenia grandifolia Cham. & Schlecht.

Millspaugh and Nuttall (1923) CA; Dunkle (1942) AN.

Franseria bipinnatifida Nutt.

Ambrosia camphorata (Greene) Payne
Watson (1876) GU.

Ambrosia chamissonis (Less.) Greene

Greene (1887a) CR; Greene (1887b) MI; Yates (1889)
MI, CR; Brandegee ( 1 890a) CA; Brandegee ( 1 890b)
MI, CR, CA; Hoffmann (1932b) RO, AN; Munz
(1935) on the islands; Eastwood (1941) MI, RO,
CR, AN, CA, CL; Dunkle (1942) AN.

Franseria bipinnatifida Nutt. var. dubia Eastw.

Ambrosia chamissonis (Less.) Greene

Eastwood (1898) NI; Eastwood (1941) NI.

Franseria camphorata Greene

Ambrosia camphorata (Greene) Payne

Greene (1885) based on Greene s.n. Apr. 24, 1 885 CAS!
GU; Vasey and Rose (1890) GU; Eastwood (1929)
GU; Howell (1942) GU.

Franseria chamissonis Less.

Ambrosia chamissonis (Less.) Greene

Greene ( 1887b) MI; Yates ( 1 889) MI; Brandegee ( 1 890b)
MI; Hoffmann (1932b) MI, CR; Munz (1935) MI,
CR, CL; Eastwood (1941) MI, CR; Munz and Keck
(1959) MI, CR, CL.

Franseria chamissonis Less. ssp. bipinnatisecta (Less.) Wig-
gins & Stockw.

Ambrosia chamissonis (Less.) Greene

Munz and Keck (1959) Channel Islands; Raven (1963)
CL.

Franseria chamissonis Less. var. viscida Eastw.

Ambrosia chamissonis (Less.) Greene

Eastwood (1898) based on Trask 10 [sic] CAS!; East-
wood (1941) NI.

Gaertneria bipinnatifida Kuntze
Ambrosia chamissonis (Less.) Greene
Millspaugh and Nuttall (1923) CA.

Gaertneria chamissonis Kuntze
Ambrosia chamissonis (Less.) Greene
Davidson and Moxley (1923) MI.

Galium angustifolium Nutt, ex T. & G.

Galium angustifolium Nutt, ex T. & G. ssp. foliosum (Hil-
lend & Howell) Dempst. & Steb.

Greene (1887a) CR; Brandegee (1888) RO; Yates (1889)
RO, CR; Brandegee (1890b) RO, CR.

Galium angustifolium Nutt, ex T. & G. var. foliosum Hillend
& Howell

omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Munz and Keck (1959) BA.

probably Galium angustifolium Nutt, ex T. & G. ssp. an-
gustifolium
Eastwood (1941) CA.

Galium aparine L. var. vaillantii (DC.) Koch
Galium aparine L.

Lyon (1886) CA; Davidson (1896) CA; Brandegee
(1890b) RO, CR, CA, CL.

Galium buxifolium Greene
omitted as unsubstantiated on CA.

Davidson (1896) CA.

Galium californicum H. & A.
omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Eastwood (1941) CL.

Galium californicum H. & A. ssp. flaccidum (Greene)
Dempst.

Abrams and Ferris (1923-1960) CR; Eastwood (1941)
CR.

Galium californicum H. & A. var. californicum

Galium californicum H. & A. ssp. flaccidum (Greene)
Dempst.

Munz and Keck (1959) CR.

Galium catalinense Gray
Galium buxifolium Greene
Brandegee (1890b) MI, CR; Dunkle (1950) MI, CR.
Galium catalinense Gray ssp. acrispum Dempst.

Abrams and Ferris (1923-1960) CL; Dunkle (1950) CL;
Munz and Keck (1959) CL.

Galium catalinense Gray var. catalinense
Galium catalinense Gray ssp. acrispum Dempst.

Raven (1963) CL.

Galium flaccidum Greene

Galium californicum H. & A. ssp. flaccidum (Greene)
Dempst.

Greene (1887a) CR; Yates (1889) CR; Eastwood (1941)
CR.

108 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Galium miguelense Greene
omitted as unsubstantiated on CA.

Davidson (1896) CA.

Galium nuttallii Gray

Galium californicum H. & A. ssp. miguelense (Greene)
Dempst. & Steb.

Brandegee (1890b) MI.

Galium nuttallii Gray var. nuttallii
Galium nuttallii gray ssp. insulare Ferris
Thome (1967) CA.

Galium nuttallii Gray ssp. ovalifolium (Dempst.) Dempst. &
Steb.

Gallium porrigens Dempst. var. porrigens (accd. Demps-
ter, 1974)

Munz (1974) CR, CA.

Galium siccatum Wight

Galium angustifolium Nutt, ex T. & G. ssp. angustifolium
Millspaugh and Nuttall (1923) CA.

Galium angustifolium Nutt, ex T. & G. ssp. foliosum (Hil-
lend & Howell) Dempst. & Steb.

Eastwood (1941) RO, CR.

Galvezia juncea (Benth.) Ball
Galvezia speciosa (Nutt.) Gray
Wiggins (1980) GU.

Gasoul crystallinum (L.) Rothm.

Mesembryanthemum crystallinum L.

Smith (1976) MI, RO, CR, AN.

Gasoul nodiflorum (L.) Rothm.

Mesembryanthemum nodiflorum L.

Munz (1974) BA, CA; Smith (1976) MI, RO, CR, AN.
Gastridium australe Beauv.

Gastridium ventricosum (Gouan) Schinz. & Thell.
Davidson (1894) CA.

Genista linifolia L.

Cytisus linifolius (L.) Lam.

Millspaugh and Nuttall ( 1 923) CA; Eastwood (1941) CA.
Gilia achilleifolia Benth. ssp. ackilleifolia
omitted as unsubstantiated on CR.

Munz (1974) CR; Smith (1976) CR.

Gilia achilleifolia Benth. ssp. multicaulis (Benth.) V. Grant
& A. Grant

Gilia clivorum (Jeps.) V. Grant
Smith (1976) RO.

Gilia androsacea (Benth.) Steud.

Linanthus androsaceus (Benth.) Greene ssp. luteus (Benth.)
Mason

Brandegee (1888) RO, CR; Yates (1889) RO, CR; Bran-
degee (1890b) RO, CR; Eastwood (1941) RO, CR.
Gilia atractyloides (Benth.) Steud. var. foliaceus (Greene)
Munz

Navarretia hamata Greene var. foliacea (Greene) Thome
Munz (1935) CA.

Gilia bicolor (Nutt.) Brand
Linanthus bicolor (Nutt.) Greene ssp. bicolor

Millspaugh and Nuttall (1923) CA; Munz (1935) CA,
CL; Eastwood (1941) RO, CR, CA, CL.

Gilia divaricata Nutt.

Allophyllum gilioides (Benth.) A. Grant & V. Grant
Greene (1885) based on Greene s.n. Apr. 24, 1885 DS!,
GU.

Gilia gilioides (Benth.) Greene

omitted as unsubstantiated on CR and CL; also omitted
by Raven (1963) on CL.

Dunkle (1950) CR, CL.

Allophyllum gilioides (Benth.) A. Grant & V. Grant
Dunkle (1950) GU.

Allophyllum glutinosum (Benth.) A. Grant & V. Grant
Jepson( 1909-1 943) CA; Munz (1 935) CA; Dunkle (1950)
CA.

Gilia nevinii Gray
Dunkle (1950) BA.

Gilia gilioides (Benth.) Greene var. glutinosa (Benth.) Jeps.

omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Dunkle (1950) CL.

Allophyllum gilioides (Benth.) A. Grant & V. Grant
Dunkle (1950) GU.

Allophyllum glutinosum (Benth.) A. Grant & V. Grant
Jepson (1909-1943) based on Fosberg 15413 [sic] ac-
tually is Fosberg S5413 LAM!, POM! CA; Dunkle
(1950) CA.

Gilia nevinii Gray (Philbrick, 1972)

Dunkle (1942) BA; Dunkle (1950) BA.

Gilia glutinosa (Benth.) Gray

Allophyllum glutinosum (Benth.) A. Grant & V. Grant
Brandegee (1890a) CA; Brandegee (1890b) CA; Mill-
spaugh and Nuttall (1923) CA; Eastwood (1941)
CA.

Gilia guadalupensis Brand

probably Linanthus pygmaeus (Brand) J.T. Howell ssp.
pygmaeus

Eastwood (1929) GU.

Gilia micrantha Steud. ex Benth. in A. DC.

Linanthus bicolor (Nutt.) Greene ssp. bicolor

Lyon (1886) CL; Greene (1887b) MI; Yates (1889) MI;
Brandegee (1890b) MI, CL; Eastwood (1941) MI,
CL.

Gilia millefoliata F. & M.

Gilia clivorum (Jeps.) V. Grant
Hoffmann (1932b) MI, RO, CR, AN; Munz (1935) MI,
RO, CR, AN, CA; Eastwood (1941) MI, RO, CR,
AN, CA; Dunkle (1942) AN.

Gilia aff. multicaulis Benth.

Gilia angelensis V. Grant

Gentry (1949) based on Elmore 440 LAM!, CR (annot.
by A. Day).

Gilia multicaulis Benth.

Gilia angelensis V. Grant

Lyon (1886) CA; Greene ( 1 887a) CR; Yates (1889) CR;
Brandegee ( 1 890b) CR, CA; Millspaugh and Nuttall
(1923) CA; Abrams and Ferris (1923-1960) CR;
Eastwood (1941) CR, CA; Dunkle (1942) AN.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 109

Gilia clivorum (Jeps.) V. Grant
Jepson (1909-1943) RO, CR.

Gilia nevinii Gray

Greene (1885) based on Greene s.n. Apr. 25, 1885 CAS!,
GU.

Gilia multicaulis Benth. var. millefolia Gray
Gilia nevinii Gray

Watson (1876) based on Palmer 78 NY!, GU; Jepson
(1909-1943) RO, CR, CL based on Munz 6633
POM!, GU based on Anthony 235 GH!.

Gilia multicaulis Benth. var. nevinii (Gray) Jeps.

Gilia nevinii Gray
Jepson (1925) GU.

Gilia multicaulis Benth. var. peduncularis (Eastw.) Jeps.
Gilia nevinii Gray
Jepson (1909-1943) CR, CL, GU.

Gilia nevinii Gray
omitted as unsubstantiated on MI.

Eastwood (1941) MI.

Gilia pusilla Benth.

probably Linanthus pygmaeus (Brand) J.T. Howell ssp.
pygmaeus
Greene (1885) GU.

Gilia pusilla Benth. var. californica Gray
probably Linanthus pygmaeus (Brand) J.T. Howell ssp.
pygmaeus

Watson (1876) GU; Jepson (1909-1943) GU.

Gilia tenuiflora Benth.
omitted as unsubstantiated on CA.

Eastwood (1941) CA; Dunkle (1950) CA.

Gilia viscidula Gray

Navarretia atractyloides (Benth.) H. & A.

Brandegee (1890a) CA; Brandegee (1890b) CA.
Githopsis specularioides Nutt,
omitted as unsubstantiated on RO and CA.

Dunkle (1950) RO, CA.

Githopsis specularioides Nutt. ssp. Candida Ewan
Githopsis diffusa Gray var. guadalupensis Morin
Wiggins (1980) GU; (Morin, N. 1983. Syst. Bot. 8(4):
436-468).

Gnaphalium californicum DC.
omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Eastwood (1941) CL.

Gnaphalium chilense Spreng.

Gnaphalium bicolor Bioletti

Eastwood (1941) probably based on Abrams & Wiggins
375 DS!, misidentified, CL.

Gnaphalium chilense Spreng. var. confertifolium Greene
Gnaphalium chilense Spreng.

Eastwood (1941) MI, RO.

Gnaphalium decurrens Ives
Gnaphalium bicolor Bioletti

Lyon (1886) CL; Brandegee (1890b) CL.

Gnaphalium californicum DC.

Brandegee (1890b) RO, CR.

Gnaphalium decurrens Ives var. californicum (DC.) Gray
Gnaphalium californicum DC.

Greene (1887a) CR; Brandegee (1888) RO; Yates (1889)

RO, CR.

Gnaphalium palustre Nutt.

Filago californica Nutt.

Gentry (1949) based on Elmore 443 AHFH!, misiden-
tified, CR.

Gnaphalium sprengelii H. & A.

Gnaphalium chilense Spreng.

Watson (1876) GU; Greene (1885) GU; Greene (1887a)
CR; Greene (1887b) MI; Brandegee (1888) RO;
Yates (1889) MI, RO, CR; Brandegee (1890a) CA;
Brandegee (1890b) MI, RO, CR, CA; Vasey and
Rose (1890) based on Palmer 885 NY!, GU; Mills-
paugh and Nuttall (1923) CA; Eastwood ( 1 929) GU;
Dunkle (1950) GU.

Gnaphalium wrightii Gray

Gnaphalium microcephalum Nutt.

Hoffmann (1932b) CR; Eastwood (1941) CR.

Godetia bottae Spach.

Clarkia purpurea (Curt.) Nels. & Macbr. ssp. quadrivulnera
(Dougl. in Lindl.) Lewis & Lewis
Brandegee (1890a) CA; Brandegee (1890b) CA; East-
wood (1941) CA.

Godetia purpurea (Curt.) G. Don in Sweet
Clarkia purpurea (Curt.) Nels. & Macbr. ssp. quadrivulnera
(Dougl. in Lindl.) Lewis & Lewis
Greene (1887a) CR; Yates (1889) CR; Brandegee (1890b)
CR; Eastwood (1941) CR.

Godetia quadrivulnera (Dougl. in Lindl.) Spach. var. tenella
(Cav.) Jeps.

Clarkia purpurea (Curt.) Nels. & Macbr. ssp. quadrivulnera
(Dougl. in Lindl.) Lewis & Lewis
Eastwood (1941) CA.

Godetia tenella (Cav.) Steud.

Clarkia purpurea (Curt.) Nels. & Macbr. ssp. quadrivulnera
(Dougl. in Lindl.) Lewis & Lewis
Lyon (1886) CA; Brandegee (1890b) CA; Trask (1899)
CA.

Grindelia glutinosa Dunal
Grindelia latifolia Kell. ssp. latifolia
Brandegee (1888) RO; Brandegee (1890b) MI, RO, CR.
Grindelia latifolia Kell. ssp. platyphylla (Greene) Keck
Grindelia latifolia Keck
Smith (1976) AN.

Grindelia perennis A. Nels.
incertae sedis

Munz (1935) CA; Eastwood (1941) CA.

Grindelia robust a Nutt. var. platyphylla Greene
Grindelia latifolia Kell.

Eastwood (1941) RO, AN.

Grindelia rubricaulis DC. var. platyphylla (Greene) Steyerm.
omitted as unsubstantiated on BA.

Munz (1935) BA.

Grindelia latifolia Kell.

Munz (1935) RO, AN; Dunkle (1950) RO, AN.

110 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Habenaria elegans (Lindl.) Boland, var. maritima (Greene)
Ames

omitted as unsubstantiated on RO.

Smith (1976) RO.

Habenaria michaeli Greene
Habenaria elegans (Lindl.) Boland.

Hoffmann ( 1 932a) RO; Munz (1935) RO, CR; Eastwood
(1941) RO, CR; Dunkle (1950) RO, CR.
Haplopappus canus (Gray) Blake

Haplopappus detonsus (Greene) Raven

Abrams and Ferris (1923-1960) RO, CR; Munz and
Keck (1959) RO, CR.

Haplopappus detonsus (Greene) Raven
Haplopappus canus (Gray) Blake
Smith (1976) GU.

Haplopappus squarrosus H. & A. ssp. squarrosus
omitted as unsubstantiated on CR.

Smith (1976) CR.

Hazardia cana (Gray) Greene
Haplopappus detonsus (Greene) Raven
Davidson and Moxley (1923) RO, CR; Jepson (1925)
RO, CR; Eastwood (1941) RO, CR.

Hazardia detonsa (Greene) Greene
Haplopappus detonsus (Greene) Raven
Greene (1887a) CR; Yates (1889) CR; Ford (1890) CR;
Eastwood (1941) RO, CR; Clark (1979) RO, CR.
Hazardia serrata Greene

Haplopappus detonsus (Greene) Raven x Haplopappus
squarrosus H. & A. ssp. grindelioides (DC.) Keck
(omitted from text).

Greene (1887a) CR; Yates (1889) RO, CR; Ford (1890)
CR; Eastwood (1941) RO, CR.

Hazardia squarrosa (H. & A.) Greene
Haplopappus squarrosus H. & A. ssp. grindelioides (DC.)
Keck

McClatchie (1894) CA; Millspaugh and Nuttall (1923)
CA; Jepson (1925) CR; Eastwood (1941) RO, CR.
Hazardia squarrosa (H. & A.) Greene var. grindelioides (DC.)
Clark

Haplopappus squarrosus H. & A. ssp. grindelioides (DC.)
Keck

Clark (1979) MI, RO, CR, AN, CA.

Heleniastrum puberulum (DC.) Kuntze
Helenium puberulum DC.

McClatchie (1894) CA.

Helianthemum scoparium Nutt. var. vulgare Jeps.
Helianthemum scoparium Nutt.

Munz and Keck (1959) CA; Thome (1967) CA; Smith
(1976) RO“?,” CR.

Heliotropium chenopodioides Willd.

Heliotropium curassavicum L. ssp. oculatum (Heller)
Thome

Millspaugh and Nuttall (1923) CA.

Hemizonia fasciculata (DC.) T. & G. var. ramosissima

(Benth.) Gray

Hemizonia fasciculata (DC.) T. & G.

Eastwood (1941) BA, CA, CL; Dunkle (1950) BA, CA,
CL.

Hemizonia floribunda Gray
Hemizonia Clementina Bdg.

Millspaugh and Nuttall (1923) based in part on Nuttall
195 F! and Nuttall 352 F!, misidentified, CA; East-
wood (1941) CA.

Hemizonia paniculata Gray
Hemizonia Clementina Bdg.

Millspaugh and Nuttall (1923) based in part on Knopf
148 US! and Smith 5055 US!, misidentified, CA;
Eastwood (1941) CA.

Hemizonia paniculata Gray

Hemizonia increscens (Hall ex Keck) Tanowitz spp. in-
crescens

Brandegee (1888) RO; Yates (1889) RO; Brandegee
(1890b) RO; Hoffmann (1932b) CR; Eastwood
(1941) RO, CR.

Hemizonia paniculata Gray ssp. increscens Hall ex Keck
Hemizonia increscens (Hall ex Keck) Tanowitz ssp. in-
crescens

Abrams and Ferris (1923-1960) RO; Smith (1976) RO,
CR.

Hemizonia ramosissima Benth.

Hemizonia fasciculata (DC.) T. & G.

Smith (1976) CR.

Hemizonia streetsii Gray
omitted as unsubstantiated on GU.

Wiggins (1980) GU.

Hemizonia Clementina Bdg.

Lyon (1886) CA; Davidson (1896) based on Davidson
s.n. LAM! CA; Yates ( 1 889) AN; Brandegee ( 1 890b)
AN, CA, CL; Eastwood (1898) NI, CA, CL.
Hemizonia wrightii Gray
Hemizonia fasciculata (DC.) T. & G.

Brandegee (1890b) CA; Davidson (1896) CA.
Hesperastragalus didymocarpus (H. & A.) Heller
Astragalus didymocarpus H. & A.

Millspaugh and Nuttall (1923) CA.

Hesperastragalus gambelianus (Sheld.) Heller
Astragalus gambelianus Sheld.

Millspaugh and Nuttall (1923) CA.

Hesperonia californica (Gray) Standi.

Mirabilis californica Gray var. californica
Millspaugh and Nuttall (1923) CA.

Hesperonia californica (Gray) Standi, var. microphylla Standi.
Mirabilis californica Gray var. californica
Eastwood (1941) RO, CR, CA, CL.

Hesperonia cendrosensis Standi.

Mirabilis californica Gray var. californica

Jepson (1909-1943) (doubtfully attributable to CL); Da-
vidson and Moxley (1923) CL; Eastwood (1941) CL.
Hesperonia heimerlii Standi.

Mirabilis heimerlii (Standi.) Macbr.

Eastwood (1929) GU.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 111

Hesperonia laevis (Benth.) Standi.

Mirabilis californica Gray var. californica
Eastwood (1941) RO; Gentry (1949) CL.

Heuchera sp.

possibly Jepsonia malvaefolia (Greene) Small
Watson (1876) based on Palmer s.n. in 1876 GH! sterile;
Franceschi (1893) GU; Eastwood (1929) GU.
Heuchera micrantha Dougl.

Heuchera maxima Greene
Clokey (1931) based on Clokey 4952 LAM!, CR.
Heuchera pilosissima F. & M.

Heuchera maxima Greene

Brandegee (1888) RO; Brandegee (1890b) RO, CR.
Hieraceum argutum Nutt.

omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Eastwood (1941) BA.

Hieraceum grinellii Eastw.

Hieraceum arguta Nutt.

Clokey (1931) CR; Eastwood (1941) CR.

Hirschfeldia incana (L.) Lagr.-Foss.

Brassica geniculata (Desf.) Ball.

Raven (1963) CL; Foreman (1967) NI.

Holodiscus ariaefolius (Sm. in Rees) Greene

Holodiscus discolor (Pursh) Maxim, ssp. discolor
Eastwood (1941) CR, CA.

Holodiscus discolor (Pursh) Maxim, var. franciscanus (Rydb.)
Jeps.

Holodiscus discolor (Pursh) Maxim, ssp. discolor

Jepson (1909-1943) CR; Thome (1967) based on Thome
36905 RSA! and Fosberg S5436 POM!, CA; Smith
(1976) CR.

Hookera minor Britten
Brodiaea jolonensis Eastw.

Millspaugh and Nuttall (1923) CA.

Hordeum glaucum Steud.

Hordeum murinum L. ssp. glaucum (Steud.) Tzvel.

Foreman (1967) NI; Smith (1976) all four islands.
Hordeum murinum L. ssp. leporinum (Link) Arcang.
Foreman (1967) based on Foreman & Lloyd 132 LA!,
misidentified, NI.

Hordeum gussoneanum Pari.

Hordeum geniculatum All.

Hoffmann (1932a) RO; Eastwood (1941) RO.

Hordeum hystrix Roth.

Hordeum geniculatum All.

Thome (1967) CA.

Hordeum leporinum Link

Hordeum murinum L. ssp. leporinum (Link) Arcang.
Foreman (1967) NI; Smith (1976) all four islands.
Hordeum murinum L.
incertae sedis

Greene (1885) GU; Lyon (1886) CA; Greene (1887a)
CR; Brandegee (1888) RO; Yates (1889) RO, CR,
AN; Brandegee (1890b) RO, CR, AN; Hoffmann
(1932a) MI; Howell (1935) NI; Eastwood (1941)
MI, RO, CR, NI, CA.

Hordeum murinum L. ssp. glaucum (Steud.) Tzvel.

Vasey and Rose (1890) based on Palmer 671 F!, GU;
Dunkle (1942) based on Dunkle 8108 LAM!, AN.
Hordeum murinum L. ssp. leporinum (Link) Arcang.
Eastwood (1929) based on Mason 1544 F!, GU.
Hordeum nodosum L.

possibly Hordeum californicum Covas & Steb.

Dunkle (1942) AN.

probably Hordeum geniculatum All. (Raven 1963).

Lyon (1 886) CL; Brandegee ( 1 890b) CL; Millspaugh and
Nuttall (1923) CA.

Hordeum stebbinsii Covas
Hordeum murinum L. ssp. glaucum (Steud.) Tzvel.
Wiggins (1980) GU.

Hosackia anthylloides (Gray) Millsp. & Nutt.

Lotus grandiflorus (Benth.) Greene var. grandiflorus
Millspaugh and Nuttall (1923) CA; Eastwood (1941) CA.
Hosackia argophylla Gray
incertae sedis

Brandegee (1890b) CR, CL.

Lotus argophyllus (Gray) Greene ssp. ornithopus (Greene)
Raven

Watson (1876) GU; Brandegee (1890b) CA; Franceschi
(1893) GU; Trask (1899) CA.

Hosackia dendroidea (Greene) Abrams
Lotus scoparius (Nutt, in T. & G.) Ottley var. dendroideus
(Greene) Ottley

Abrams and Ferris (1923-1960) RO, CR, AN, CA.
Hosackia glabra (Vog.) Torr.

Lotus scoparius (Nutt, in T. & G.) Ottley ssp. scoparius
Lyon (1886) CA; Brandegee (1888) RO; Brandegee
(1890b) MI, RO, CR, CA.

Hosackia nivea (Greene) Wats.

Lotus argophyllus (Gray) Greene ssp. niveus (Greene) Munz
Abrams and Ferris (1923-1960) CR.

Hosackia occulta Greene

Lotus grandiflorus (Benth.) Greene var. grandiflorus
Greene (1887a) CR; Yates (1889) CR; Brandegee ( 1 890b)
CR.

Hosackia ornithopus Greene

omitted as unsubstantiated on MI.

Abrams and Ferris (1923-1960) MI.

Hosackia ornithopus Greene ssp. venusta (Eastw.) Abrams
Lotus argophyllus (Gray) Greene ssp. ornithopus (Greene)
Raven

Abrams and Ferris (1923-1960) NI, CA.

Hosackia purshiana Benth.

Lotus purshianus (Benth.) Clem. & Clem. var. purshianus
Greene (1887a) CR; Yates ( 1 889) CR; Brandegee ( 1 890a)
CA; Brandegee (1890b) CR, CA.

Hosackia wrangeliana (F. & M.) T. & G.
omitted as unsubstantiated on CL.

Eastwood (1941) CL.

Lotus subpinnatus Lag.

Millspaugh and Nuttall (1923) CA; Eastwood (1941) CR,
CA.

112 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Ipomoea hederacea (L.) Jacq.

Ipomoea nil (L.) Roth

Jepson ( 1 909-1943) CA; Millspaugh and Nuttall ( 1 923)
CA; Eastwood (1941) CA; Thome (1967) CA.

Isocoma latifolia Greene

Haplopappus venetus (HBK.) Blake ssp. vernonioides (Nutt.)

Hall

Eastwood (1941) RO, CR.

Isocoma veneta (HBK.) Greene var. decumbens (Bdg.) Jeps.

Haplopappus venetus (HBK.) Blake ssp. vernonioides (Nutt.)

Hall

Hoffmann ( 1 932b) based on Hoffmann s.n. SBM#209 1 !,
misidentified, RO, based on Hoffmann s.n.
SBM#10278!, misidentified, AN.

Isomeris arborea Nutt.

Cleome isomeris Greene

Lyon (1886) CA; Brandegee (1888) RO; Yates (1889)
RO; Brandagee (1890b) RO, CA; Millspaugh and
Nuttall (1923) CA; Munz (1935) CA.

Isomeris arborea Nutt. var. globosa Cov.

Cleome isomeris Greene

Jepson (1909-1943) CA, RO; Munz (1935) RO, CA;
Eastwood (1941) RO, CR, CA; Dunkle (1950) RO,
CR, CA.

Isomeris arborea Nutt. var. insularis Jeps.

Cleome isomeris Greene

Jepson ( 1 909-1943) RO, CA; Eastwood (1941) RO, CA;
Munz and Keck (1959) RO, CA; Munz (1974) RO,
CA; Smith (1976) RO, CA.

Jepsonia neo-nuttalliana Millsp. in Millsp. & Nutt.

Jepsonia malvaefolia (Greene) Small

Millspaugh and N uttall (1923) CA; Eastwood (1941) CA.

Jepsonia parryi (Torn.) Small

Jepsonia malvaefolia (Greene) Small

Jepson (1909-1943) RO, CR, CA, GU; Jepson (1925)
RO, CR; Howell (1935) NI; Munz and Keck (1959)
Channel Islands; Foreman (1967) NI.

Juncus bufonius L. var. congestus Wahlb.

Juncus bufonius L.

Wiggins (1980) GU.

Juncus bufonius L. var. halophilus Buch. & Fem.

Juncus bufonius L.

Jepson (1909-1943) CR.

Juncus effusus L.

incertae sedis

Greene (1887a) CR; Yates (1889) CR; Brandegee ( 1 890b)
CR.

Juncus robust us Wats.

Juncus acutus L. var. sphaerocarpus Engelm.

Brandegee (1890a) CA; Brandegee (1890b) CA; Mill-
spaugh and Nuttall (1923) CA; Eastwood (1941)
CA.

Koeleria cristata (L.) Pers.

omitted as unsubstantiated on CA and CL.

Eastwood (1941) CA, CL.

Koeleria pyramidata (Lam.) Beauv.

Brandegee ( 1 888) RO, CR; Yates ( 1 889) RO, CR; Bran-
degee ( 1 890b) RO, CR; Munz (1935) RO, CR; East-
wood (1941) RO, CR.

Koeleria macrantha (Ledeb.) Spreng.

Koeleria pyramidata (Lam.) Beauv.

Smith (1976) RO, CR.

Krynitzkia ambigua Gray
incertae sedis

Lyon (1886) CA, CL; Brandegee (1890b) CA, CL.

Krynitzkia foliosa Greene

Cryptantha foliosa (Greene) Greene
Greene (1885) GU; Franceschi (1893) GU; Vasey and
Rose (1890) GU.

Krynitzkia intermedia Gray

Cryptantha intermedia (Gray) Greene
Brandegee (1890a) CA; Brandegee (1890b) CA.

Krynitzkia leiocarpa F. & M.

Cryptantha leiocarpa (F. & M.) Greene
Greene (1887a) CR; Greene (1887b) MI; Brandegee
(1888) RO; Yates (1889) MI, RO, CR; Brandegee
(1890b) MI, RO, CR.

Krynitzkia maritima Greene

Cryptantha maritima (Greene) Greene
Greene (1885) GU; Franceschi (1893) GU; Vasey and
Rose (1890) GU.

Krynitzkia micromeres Gray

Cryptantha micromeres (Gray) Greene
Greene (1887a) CR; Yates (1889) CR; Brandegee (1890b)
CR.

Krynitzkia ramosissima Greene

Cryptantha maritima (Greene) Greene
Lyon (1886) CA; Davidson (1896) CA; Brandegee
(1890b) CA.

Lactuca virosa L.

Lactuca serriola L.

Millspaugh and Nuttall (1923) CA; Eastwood (1941) RO,
CR, CA.

Lastarriaea chilensis Remy
Chorizanthe coriacea Goodm.

Brandegee (1890a) CA; Brandegee (1890b) CA; David-
son (1896) CA; Millspaugh and Nuttall (1923) CA;
Hoffmann (1932a) RO, CR; Eastwood (1941) CR,
CA.

Lasthenia chrysostoma (F. & M.) Greene
Lasthenia californica DC. ex Lindl. (Johnson & Omduff
1978)

Raven (1963) CL; Thome (1967) CA; Philbrick (1972)
BA; Smith (1976) all four islands; Wiggins (1980)
GU.

Lasthenia hirsutula Greene
Lasthenia californica DC. ex Lindl.

Hoffmann (1932b) MI, RO, CR.

Lasthenia macrantha (Gray) Greene
Lasthenia californica DC. ex Lindl.

Smith (1976) MI, RO.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 113

Lathyrus alefeldii White

omitted as unsubstantiated on RO, CR, and CL.

Abrams and Ferris (1923-1960) CL; Eastwood (1941)
RO, CR; Wiggins (1980) CL.

Lathyrus strict us Nutt.
incertae sedis

Hoffmann ( 1932b) RO; Munz (1935) CA; Dunkle ( 1 950)
RO, CR, CA.

Lathyrus laetiflorus Greene ssp. barbarae (White) C. L.
Hitchc.

Munz (1935) CL; Dunkle (1950) CL.

Lathyrus vestitus Nutt, ex T. & G.
probably Lathyrus laetiflorus Greene ssp. alefeldii (White)
Brads.

Lyon (1886) CA; Brandegee (1890b) CA.

Lathyrus vestitus Nutt, ex T. & G. ssp. vestitus
Greene (1887a) CR; Yates (1889) CR; Brandegee (1890b)
CR.

Lathyrus vestitus Nutt, ex T. & G. ssp. puberulus (White ex
Greene) C.L. Hitchc.

Lathyrus vestitus Nutt, ex T. & G. ssp. vestitus
Smith (1976) FO, CR, AN.

Laurocerasus ilicifolia (Nutt, ex H. & A.) M. Roem.

Prunus lyonii (Eastw.) Sarg.

Millspaugh and Nuttall (1923) CA.

Laurocerasus lyonii (Eastw.) Britt, in Britt. & Shaf.

Prunus lyonii (Eastw.) Sarg.

Abrams (1917) islands; Davidson and Moxley (1923)
Channel Islands; Millspaugh and Nuttall (1923) CA.
Lavatera assurgentiflora Kell.

omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Raven (1967) BA.

Lavatera assurgentiflora Kell. ssp. glabra Philbrick in Power
Lavatera assurgentiflora Kell.

Philbrick (1980) CA, CL.

Layia glandulosa (Hook.) H. & A.

Layia p/atyglossa (F. & M.) Gray ssp. campestris Keck
Lyon (1886) CL.

Layia platyglossa (F. & M.) Gray
incertae sedis
Brandegee (1888) RO.

Layia platyglossa (F. & M.) Gray ssp. campestris Keck
Gentry (1949) MI, CR based on Elmore 442 AHFH!,
CA.

Layia platyglossa (F. & M.) Gray ssp. platyglossa
Gentry (1949) based on Elmore 202 AHFH!, RO.
Layia platyglossa (F. & M.) Gray ssp. platyglossa
Layia platyglossa (F. & M.) Gray ssp. campestris Keck
Abrams and Ferris (1923-1960) CR; Munz and Keck
(1959) CR; Smith (1976) CR.

Lepechinia calycina (Benth.) Epl. in Munz var. wallacei (Gray)
Epl. in Munz

Lepechinia calycina (Benth.) Epl. in Munz
Munz (1935) RO, in part.

Lepechinia fragrans (Greene) Epl.

Munz (1935) RO in part, CR, CA.

Lepidium bipinnatifidum Desv.

Lepidium oblongum Small
Eastwood (1929) GU.

Lepidium dictyotum Gray var. acutidens Gray
Lepidium latipes Hook

Hoffmann (1932b) CR; Eastwood (1941) CR, CA, CL.
Lepidium lasiophyllum Nutt.

Lepidium lasiocarpum Nutt, ex T. & G. var. lasiocarpum
Brandegee (1888) RO; Yates (1889) RO.

Lepidium medium Greene

Lepidium virginicum L. var. pubescens (Greene) Thell.
Millspaugh and Nuttall (1923) CA.

Lepidium menziesii DC.

Lepidium oblongum Small

Watson (1876) GU; Vasey and Rose (1890) based on
Palmer 897 GH!, GU.

Lepidium virginicum L. var. pubescens (Greene) Thell.
Greene (1887a) CR; Yates (1889) CR; Brandegee (1890b)
CR.

Lepidium oblongum Small var. insulare C.L. Hitchc.
Lepidium oblongum Small
Hitchcock (1945) GU.

Lepidium pubescens Desv.
incertae sedis

Eastwood (1941) AN, NI; Dunkle (1942) AN.
Lepidium oblongum Small

Hitchcock (1936) based on Trask s.n. in Mar. 1901 NY!,
misidentified, CA.

Lepidium strictum (Wats.) Rattan
Lepidium virginicum L. var. pubescens (Greene) Thell.
Eastwood ( 1 94 1 ) NI; Dunkle (1942) AN; Foreman (1967)
NI; Smith (1976) AN.

Lepigonum macrothecum F. & M.

Spergularia macrotheca (Homem.) Heynh. ssp. macro-
theca

Lyon (1886) CA; Greene (1887a) CR; Greene (1887b)
MI; Brandegee (1888) RO; Yates (1889) MI, CR;
Franceschi (1893) GU.

Lepti/on canadensis (L.) Britt, in Britt. & Br.

Conyza canadensis (L.) Cronq.

Millspaugh and Nuttall (1923) CA.

Leptilon linifolium (Willd.) Small
Conyza bonariensis (L.) Cronq.

Millspaugh and Nuttall (1923) CA.

Lepturus incurvatus (L. f.) Trin.

Parapholis incurva (L. f.) C.E. Hubb.

Jepson (1909-1943) based on Brandegee 54 US!, CA.
Lepturus paniculatus Nutt.

Parapholis incurva (L. f.) C.E. Hubb.

Brandegee ( 1 890a) probably based on Brandegee 54 US!,
CA; Brandegee (1890b) CA.

Lilium bloomerianum Kell. var. ocellatum Kell,
omitted as unsubstantiated on CA.

Eastwood (1941) CA.

Lilium humboldtii Roezl & Leichtl.

Lilium humboldtii Roezl & Leichtl. ssp. ocellatum (Kell.)
Thome

114 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Greene (1887a) CR; Brandegee (1888) RO; Yates ( 1 889)
RO, CR; Brandegee (1890b) RO, CR.

Lilium humboldtii Roezl & Leichtl. var. ocellatum (Kell.)
Elwes

omitted as unsubstantiated on CA.

Eastwood (1941) CA.

Lilium humboldtii Roezl & Leichtl. ssp. ocellatum (Kell.)
Thome

Jepson (1909-1943) RO; Jepson (1925) RO; Munz (1935)
RO, CR; Smith (1976) RO, CR.

Limonium californicum (Boiss.) Heller

probably Limonium perezii (Stapf.) F.T. Hubb.

Abrams and Ferris (1923-1960) CL.

Linanthus sp.
incertae sedis
Howell (1942) GU.

Linanthus parviflorus Greene
incertae sedis
Jepson (1909-1943) CR.

Linaria texana Sheele

Linaria canadensis (L.) Dum.-Cours. var. texana (Sheele)
Penn.

Wiggins (1980) GU.

Lithophragma catalinae Rydb.

Lithophragma affine Gray ssp. mixtum R.L. Taylor
Davidson and Moxley (1923) CA; Millspaugh and Nut-
tall (1923) CA; Eastwood (1941) CA.
Lithophragma cymbalaria T. & G.
omitted as unsubstantiated on CA; also omitted by Thome
(1967).

Eastwood (1941) CA.

Lolium multiflorum Lam.

Lolium perenne L. ssp. multiflorum (Lam.) Husnot
Raven (1963) CL; Thorne (1967) CA; Foreman (1967)
NI.

Lolium temulentum L. var. arvense (With.) Bab.

Lolium temulentum L.

Hoffmann (1932a) CR.

Lolium temulentum L. var. leptochaeton A. Br.

Lolium temulentum L.

Smith (1976) CR.

Lonicera hispidula (Lindl.) Dougl. ex T. & G. var. subspicata
(H. & A.) Gray

Lonicera subspicata H. & A. var. johnstonii Keck

Ford (1890) CR; Brandegee (1890a) CA; Brandegee
(1890b) CR, CA; Eastwood (1941) CR.

Lotus argophyllus (Gray) Greene var. niveus (Greene) Ottley
Lotus argophyllus (Gray) Greene ssp. adsurgens (Dunkle)
Raven

Ottley (1923) based on Brandegee s.n. Aug. 25, 1894
CL; Jepson (1925) CL.

Lotus argophyllus (Gray) Greene var. ornithopus (Greene)
Ottley

omitted as unsubstantiated on CR.

Dunkle (1950) CR.

Lotus argophyllus (Gray) Greene ssp. adsurgens (Dunkle)
Raven

Munz and Keck (1959) CL.

Lotus dendroideus (Greene) Greene

Lotus scoparius Nutt, in T. & G. var. dendroideus (Greene)
Ottley

Davidson (1896) CA; Gentry (1949) RO, CR, AN, CA.
Lotus dendroideus (Greene) Greene var. traskiae (Eastw. ex
Noddin in Abrams) Isely

Lotus scoparius (Nutt, in T. & G.) Ottley ssp. traskiae
(Eastw. ex Noddin in Abrams) Raven
Isely (1978) CL.

Lotus dendroideus (Greene) Greene var. veatchii (Greene)
Isely

Lotus scoparius (Nutt, in T. & G.) Ottley var. veatchii
(Greene) Ottley
Isely (1978) MI.

Lotus eriophorus Greene var. heermannii (Dur. & Hilg.) Ot-
tley

Lotus heermannii (Dur. & Hilg.) Greene
Ottley (1923) based on Grant 716 CA.

Lotus grandiflorus (Benth.) Greene var. mutabilis Ottley
Lotus grandiflorus (Benth.) Greene var. grandiflorus
Clokey (1931) CR.

Lotus micranthus Benth.

probably Lotus hamatus Greene
Davidson (1896) CA.

Lotus niveus (Greene) Greene

Lotus argophyllus (Gray) Greene ssp. adsurgens (Dunkle)
Raven

Gentry (1949) based on Elmore 410 AHFH!, misiden-
tihed, CL.

Lotus argophyllus (Gray) Greene ssp. niveus (Greene) Munz
Gentry (1949) CR.

Lotus argophyllus (Gray) Greene ssp. ornithopus (Greene)
Raven

Gentry (1949) based on Elmore 409 AHFH!, misiden-
tified, CL.

Lotus scoparius (Nutt, in T. & G.) Ottley var. dendroideus
(Greene) Ottley

Lotus scoparius (Nutt, in T. & G.) Ottley ssp. traskiae
(Eastw. ex Noddin in Abrams) Raven
Munz and Keck (1959) CL in part.

Lotus scoparius ( Nutt, in T. & G.) Ottley var. traskiae (Eastw.
ex Noddin in Abrams) Ottley
Lotus scoparius (Nutt, in T. & G.) Ottley var. dendroideus
(Greene) Ottley

Dunkle (1950) probably based on Dunkle 2056 AHFH!,
misidentified, CA.

Lupinus affinis J. G. Agardh

Lupinus succulentus Dougl. ex Koch

Lyon (1886) CL; Greene (1887a) CR; Yates (1889) CR;
Brandegee (1890b) CR; Millspaugh and Nuttall
(1923) CA.

Lupinus albifrons Benth.

omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Eastwood (1941) CL.

Lupinus albifrons Benth. var. eminens (Greene) C.P. Sm.
probably Lupinus albifrons Benth. ssp. albifrons
Hoffmann (1932b) CR; Smith (1976) CR.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 115

Lupinus arboreus Sims

omitted as unsubstantiated on CA.

Eastwood (1941) CA.

Lupinus bicolor Lindl. var. rostratus (Eastw.) Jeps.
incertae sedis
Jepson (1909-1943) CR.

Lupinus bicolor Lindl. ssp. trifidus (Torr.) C.P. Sm.
omitted as unsubstantiated on CR.

Smith (1976) CR.

Lupinus chamissonis Esch.
incertae sedis

Greene (1887a) CR; Greene (1887b) MI; Brandegee
(1888) RO; Yates (1889) MI, RO, CR; Eastwood
(1941) MI, RO, CR; Smith (1976) MI.

Lupinus clementinus Greene
Lupinus guadalupensis Greene

Davidson and Moxley (1923) CA; Dunkle (1950) CL.
Lupinus concinnus J.G. Agardh var. agardhianus (Heller)
C.P. Sm.

Lupinus agardhianus Heller
Hoffmann (1932b) RO, CR; Munz (1935) on the islands;
Raven (1963) CL.

Lupinus excubitus Jones var. hallii (Abrams) C.P. Sm.
Lupinus albifrons Benth. ssp. albifrons
Eastwood (1941) NI, CA.

Lupinus gracilis J.G. Agardh
Lupinus agardhianus Heller

Millspaugh and Nuttall ( 1 923) CA; Eastwood ( 1 94 1) CA.
Lupinus micranthus Dougl. in Lindl.
incertae sedis

Brandegee (1888) RO; Brandegee (1890b) RO.

Lupinus bicolor Lindl. ssp. microphyllus (Wats.) D. Dunn
Brandegee (1890a) CA; Brandegee (1890b) CR, CA;
Eastwood (1898) NI; Millspaugh and Nuttall (1923)
CA.

Lupinus microcarpus Sims
incertae sedis

Greene (1887a) CR; Yates (1889) CR; Brandegee (1890b)
CR; Eastwood (1941) CR.

Lupinus microcarpus Sims var. horizontalis Jeps.
incertae sedis
Jepson (1909-1943) CR.

Lupinus microcarpus Sims var. insularis C.P. Sm.
incertae sedis
Jepson (1909-1943) CR.

Lupinus moranii Dunkle

Lupinus guadalupensis Greene

Dunkle (1943) based on Moran 587 LAM!, MO!, NY!,
RSA! CL; Dunkle (1950) CL; Munz and Keck (1959)
CL.

Lupinus nanus Dougl. in Benth.

Lupinus guadalupensis Greene

Dunkle (1950) based on Moran 587 LAM!, MO!, NY!,
RSA!, misidentified, CL.

Lupinus sparsiflorus Benth.

Lupinus bicolor Lindl. ssp. microphyllus (Wats.) D. Dunn
Gentry (1949) based on Elmore 446 AHFH!, misiden-
tified, CR; Smith (1976) CR.

Lupinus sparsiflorus Benth. var. pondii (Greene) C.P. Sm.
Lupinus guadalupensis Greene
Wiggins (1980) GU.

Lupinus subvexus C.P. Sm. var. phoeniceus C.P. Sm.
incertae sedis
Smith (1976) RO, CR.

Lupinus variicolor Steud.

Lupinus arboreus Sims

Smith (1976) based on Blakley 5 1 04 SBBG! and Blakley
5854 SBBG!, misidentified, MI.

Luzula campestris (L.) DC. var. congesta (Thuill.) Meyer
Luzula subsessilis (Wats.) Buch.

Hoffmann (1932a) RO, CR; Munz (1935) RO, CR; East-
wood (1941) RO, CR; Dunkle (1950) RO, CR.

Luzula comosa Meyer
Luzula subsessilis (Wats.) Buch.

Brandegee (1888) RO, CR; Yates (1889) RO, CR; Bran-
degee (1890b) RO, CR; Eastwood (1941) MI, RO,
CR.

Lycium brevipes Benth. var. hassei (Greene) C.L. Hitchc.
Lycium verrucosum Eastw.

Jepson (1909-1943) NI.

Lycium californicum Nutt.

omitted as unsubstantiated on MI.

Dunkle (1950) MI.

Lycium richii Gray

Lycium brevipes Benth. var. hassei (Greene) C.L. Hitchc.
Brandegee (1890a) CA; Brandegee (1890b) CA; David-
son (1896) CA; Trask (1899) CA; Abrams (1917)
CA; Davidson and Moxley (1923) CA; Jepson (1925)
CA; Eastwood (1941) CA, CL; Dunkle (1950) CA.
Lycium richii Gray var. hassei (Greene) Jtn.
omitted as unsubstantiated on GU.

Dunkle (1950) GU.

Mahonia pinnata (Lag.) Fedde.

Berberis pinnata Lag. ssp. insularis Munz
Eastwood (1941) RO, CR.

Malacothrix blairii (M. & J.) Munz
Stephanomeria blairii M. & J.

Abrams and Ferris (1923-1960) CL; Munz (1935) CL;
Eastwood ( 1 94 1 ) CL; Dunkle ( 1 9 50) CL; Munz and
Keck (1959) CL.

Malacothrix californica DC.

omitted as unsubstantiated on CR and CA.

Munz (1935) CR, CA.

Malacothrix clevelandii Gray
Malacothrix" A” (Davis, 1980)

Gentry (1949) based on Elmore 370 AHFH! BA.
Malacothrix" C” (Davis, 1980)

Foreman (1967) based on Foreman, Evans and Rainey
80 UC! NI.

Malacothrix similis Davis & Raven

Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890b) CR; Hall (1907) CR; Abrams and Ferris
(1923-1960) CR; Munz (1935) CR; Eastwood (1941)
CR; Gentry (1949) CR; Smith (1976) CR.

116 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Malacothrix squalida Greene

Eastwood (1941) AN; Dunkle ( 1 942) AN; Gentry ( 1 949)
AN; Smith (1976) AN.

Malacothrix coulteri Harv. & Gray var. coulteri

Malacothrix coulteri Harv. & Gray var. cognata Jeps.
Williams (1957) RO, CR.

Malacothrix foliosa Gray
incertae sedis

Brandegee (1890b) MI, CR; Jepson (1925) CR.
omitted as column transposition for BA.

Dunkle (1950) CA.

Malacothrix incana (Nutt.) T. & G. var. incana
Gentry (1949) based on Elmore 338 AHFH!, misiden-
tified, MI.

probably Malacothrix indecora Greene.

Brandegee (1890b) MI, CR; Williams (1957) based on
Brandegee s.n. in 1888 at UC. not seen, CR.

Malacothrix foliosa Gray var. indecora (Greene) Williams
incertae sedis
Williams (1957) NI.

Malacothrix indecora Greene
Williams (1957) MI, CR.

Malacothrix foliosa Gray var. squalida (Greene) Williams
incertae sedis
Williams (1957) MI.

probably Malacothrix “A” (Davis, 1980).

Williams (1957) BA.

Malacothrix squalida Greene
Williams (1957) CR.

Malacothrix incana (Nutt.) T. & G. var. succulenta (Elmer)
Williams

Malacothrix incana (Nutt.) T. & G.

Smith (1976) MI, RO, CR.

Malacothrix indecora Greene
Malacothrix “C” (Davis, 1980)

Eastwood (1898) NI; Jepson (1925) NI.

Malacothrix insularis Greene
incertae sedis

Brandegee (1890b) CR; Eastwood (1941) CR.

Malacothrix insularis Greene var. squalida (Greene) Ferris
incertae sedis

Abrams and Ferris (1923-1960) RO.

Malacothrix “A” (Davis, 1 980)

Abrams and Ferris (1923-1960) BA.

Malacothrix squalida Greene
Abrams and Ferris (1923-1960) CR, AN.

Malacothrix saxatitis (Nutt.) T. & G.

Malacothrix saxatilis { Nutt.) T. & G. var. implicata (Eastw.)
Hall

Brandegee (1888) RO; Yates (1889) RO; Brandegee
(1890b) MI, RO, CR.

Malacothrix saxatilis (Nutt.) T. & G. var. tenuifolia (Nutt.)
Gray

Lyon (1886) CA; Brandegee (1890b) CA; Davidson
(1896) CA; Jepson (1925) CA.

Malacothrix similis Davis & Raven
Malacothrix squalida Greene (according to Davis pers.
comm.).

Smith (1976) AN.

Malacothrix squalida Greene

omitted as unsubstantiated on RO.

Abrams and Ferris (1923-1960) RO; Munz and Keck
(1959) RO; Smith (1976) RO.

Malacothrix “A” (Davis, 1980)

Abrams and Ferris (1923-1960) BA; Munz and Keck
(1959) BA.

probably Malacothrix indecora Greene.

Eastwood (1941) MI.

Malacothrix tenuifolia T. & G.

Malacothrix saxatilis (Nutt.) T. & G. var. implicata (Eastw.)
Hall

Greene (1887a) CR; Greene (1887b) MI; Yates (1889)
MI, CR, AN.

Malva borealis Wallr.

Malva parviflora L.

Watson (1876) GU; Greene (1885) GU; Lyon (1886)
CL; Brandegee (1888) RO; Brandegee (1890a) CA;
Brandegee (1890b) MI, RO, CR, CA, CL; Vasey
and Rose (1890) GU; Franceschi (1893) GU; East-
wood (1929) GU.

Malva nicaensis All.

Malva parviflora L.

Smith (1976) CR.

Malva pusilta Sm. in Sowerby
Malva parviflora L.

Eastwood (1898) NI; Eastwood (1941) MI, RO, CR, NI,
CA.

Malvastrum exile Gray

Eremalche exilis (Gray) Greene

Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890b) CR; Ford (1890) CR; Trask (1904) CL;
Eastwood (1941) CR, CL; Thome (1967) CA.

Malvastrum fasciculatum Greene
Malacothamnus fasciculatus (Nutt.) Greene ssp. catali-
nensis (Eastw.) Thome

Millspaugh and Nuttall (1923) CA; Dunkle (1950) CA.
Malacothamnus fasciculatus (Nutt.) Greene var. nesioticus
(Rob.) Keam.

Dunkle (1950) CR.

Malvastrum thurberi Gray

Malacothamnus fasciculatus (Nutt.) Greene ssp. catali-
nensis (Eastw.) Thome

Lyon (1886) CA; Brandegee (1890b) CA; Trask (1899)
CA.

Malacothamnus fasciculatus (Nutt.) Greene var. nesioticus
(Rob.) Keam.

Brandegee (1890b) CR.

Malvastrum thurberi Gray var. laxiflorum Gray
Malacothamnus fasciculatus (Nutt.) Greene var. nesioticus
(Rob.) Keam.

Greene (1887a) CR; Yates (1889) CR; Ford (1890) CR.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 117

Mammillaria goodridgei Scheer in Salm-Dyck

Mammillaria blossfeldiana Boedeker var. shurliana Gates
Greene (1885) GU; Vasey and Rose (1890) GU; Howell
(1942) GU.

Marah fabacea (Naud.) Greene
Marah macrocarpus (Greene) Greene
Gentry (1949) CL; Smith (1976) RO.

Marah guadalupensis (Wats.) Greene
Marah macrocarpus (Greene) Greene
Abrams and Ferris (1923-1960) NI, CA, CL; Eastwood
(1941) MI, RO, CR.

Marah macrocarpus (Greene) Greene var. major (S.T. Dunn)
Stocking

Marah macrocarpus (Greene) Greene
Munz and Keck (1959) Channel Islands.

Marah major S.T. Dunn
Marah macrocarpus (Greene) Greene

Howell (1935) NI; Eastwood (1941) NI, CA, CL.
Marrubium vulgare L.

omitted as unsubstantiated on BA.

Gentry (1949) BA.

Maruta cotula (L.) DC.

Anthemis cotula L.

Millspaugh and Nuttall (1923) CA.

Meconel/a oregana Nutt, in T. & G. var. denticulata (Greene)
Jeps.

Meconella denticulata Greene
Jepson (1909-1943) CR; Munz and Keck (1959) CR.

Medicago apiculata Willd.

Medicago polymorpha L. var. brevispina (Benth.) Heyn.
Jepson (1909-1943) CR; Clokey (1931) CR; Hoffmann
(1932b) RO, CR; Eastwood (1941) RO, CR, CA,
CL.

Medicago denticulata Willd.

Medicago sativa L.

Lyon (1886) CA, CL; Greene (1887a) CR; Greene (1 887b)
MI; Yates (1889) MI, CR; Brandegee (1890b) MI,
CR, CA, CL; Eastwood (1898) NI.

Medicago hispida Gaertn.

Medicago polymorpha L. var. polymorpha
Millspaugh and Nuttall (1923) CA; Hoffmann (1932b)
RO; Howell (1935) NI; Eastwood (1941) MI, RO,
CR, NI, CA, CL; Howell ( 1 942) GU; Dunkle ( 1 942)
AN, BA; Dunkle (1950) MI, RO, CR, AN, NI, BA,
CA, CL.

Megarrhiza californica Torr. in Wats.

Marah macrocarpus (Greene) Greene
Lyon (1886) CA.

Megarrhiza guadalupensis Wats.

Marah guadalupensis (Wats.) Greene
Watson (1876) GU.

Megarrhiza marah Wats.

Marah macrocarpus (Greene) Greene
Lyon (1886) CA.

Melica imperfecta Trin.
omitted as unsubstantiated on NI.

Dunkle (1950) NI.

Melica imperfecta Trin. var. flexuosa Boland.

Melica imperfecta Trin.

Hoffmann (1932a) CR; Munz (1935) CR; Eastwood
(1941) CR.

Melica imperfecta Trin. var. minor Scribn.

Melica imperfecta Trin.

Jepson (1909-1943) CA; Davidson and Moxley (1923)
CA; Millspaugh and Nuttall (1923) CA.

Melica imperfecta Trin. var. refracta Thurb. in Wats.
Melica imperfecta Trin.

Jepson (1909-1943) CR.

Melica torreyana Scribn.

Melica imperfecta Trin.

Millspaugh and Nuttall (1923) CA; Eastwood (1941) MI,
CR, CA.

Melilotus parviflorus Desf.

Melilotus indica (L.) All.

Lyon (1886) CA; Greene (1887a) CR; Greene (1887b)
MI; Brandegee (1888) RO; Yates (1889) MI, RO,
CR; Brandegee (1890b) MI, RO, CR, CA.

Mentha piperita L.

Mentha citrata Ehrh.

Millspaugh and Nuttall ( 1 923) CA; Eastwood ( 1 94 1) CA.
Mentzelia dispersa Wats.

Mentzelia micrantha Wats.

Watson (1876) GU; Vasey and Rose (1890) based on
Palmer 893 ND-G!, GU; Eastwood (1929) GU;
Dunkle (1950) GU.

Mentzelia gracilenta T. & G.

Mentzelia ajftnis Greene

Lyon (1886) CL; Brandegee (1890b) CL; Eastwood ( 1 94 1 )
CL.

Mentzelia gracilenta T. & G. var. eremophila Jeps.
Mentzelia affinis Greene
Jepson (1909-1943) CA.

Mesembry’anthemum aequilaterale Haw.

omitted as unsubstantiated on CA but recently reported
by Thome (pers. comm.).

Millspaugh and Nuttall (1923) CA.

Carpobrotus aequilaterus (Haw.) N.E. Br.

Greene (1887a) CR; Greene (1887b) MI; Brandegee
( 1 890b) MI, CR; Jepson ( 1 909-1943) MI, CR; Hoff-
mann (1932a) RO.

Mesembryanthemum chilense Mol.

Carpobrotus aequilaterus (Haw.) N.E. Br.

Munz (1935) MI; Eastwood (1941) MI, RO, CR, CA;
Dunkle (1942) AN; Gentry (1949) MI.
Mesembryanthemum cordifolium L. f.

Aptenia cordifolia (L. f.) N.E. Br.

Munz and Keck (1959) CA.

Mesembryanthemum edute L.

Carpobrotus edulis (L.) Bolus
Foreman (1967) NI.

Micrampelis macrocarpa (Greene) Greene
Marah macrocarpus (Greene) Greene
Millspaugh and Nuttall (1923) CA.

118 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Micropus californicus F. & M.

Filago californica Nutt.

Watson ( 1 876) based on Palmer 37 CM!, NY!, misiden-
tified, GU; Eastwood (1929) GU; Dunkle (1950)
GU.

Microseris anomala Wats.

Microseris linearifolia (DC.) Sch.-Bip. (Chambers, 1955).

Brandegee ( 1 888) CR; Yates (1889) CR; Eastwood (1941)
CR.

Microseris aphantocarpha (Gray) Sch.-Bip.

Microseris douglasii (DC.) Sch.-Bip. ssp. platycarpa (Gray)
Chamb.

Eastwood (1941) CA, CL.

Microseris lindleyi (DC.) Gray

omitted as unsubstantiated on MI.

Dunkle (1950) MI.

Microseris heterocarpa (Nutt.) Chamb.

Lyon (1886) CL; Brandegee (1888) RO, CR; Brandegee
(1890a) CA; Brandegee (1890b) RO, CR, CA, CL;
Brandegee (1900) GU; Millspaugh and Nuttall
(1923) CA; Eastwood (1929) GU; Munz (1935) on
the islands; Eastwood (1941) RO, CR, CA, CL;
Dunkle (1950) RO, CA, CL, GU.

Microseris lindleyi (DC.) Gray var. clevelandii (Greene) Hall
omitted as unsubstantiated on AN.

Dunkle (1950) AN.

Microseris heterocarpa (Nutt.) Chamb. (Chambers, 1955)

Dunkle (1950) CR, GU.

Microseris linearifolia (DC.) Sch.-Bip.
omitted as unsubstantiated on MI.

Dunkle (1950) ML

Microsteris traskiae (Eastw. ex Milliken) Davids. & Mox.
Allophyllum glutinosum (Benth.) A. Grant & V. Grant

Davidson and Moxley (1923) CA.

Mimulus douglasii Gray
Mimulus traskiae Grant in Millsp. & Nutt.

Davidson and Moxley ( 1 923) based on Trask s.n. in Mar.
1901 LAM! CA.

Mimulus flemingii Munz
Diplacus parviflorus Greene

Abrams and Ferris (1923-1960) RO, CR, AN; Munz
(1935) RO, CR; Dunkle (1942) AN; Dunkle (1950)
RO, CR, AN, CL; Munz and Keck (1959) RO, CR,
AN, CL; Raven (1963) CL; Munz (1974) RO, CR,
AN, CL.

Mimulus glutinosus Wendl.
incertae sedis

Brandegee (1890b) RO, CR.

Diplacus puniceus Nutt.

Brandegee (1890a) CA; Brandegee (1890b) CA; Trask
(1899) CA.

Mimulus glutinosus Wendl. var. puniceus (Nutt.) Gray
Diplacus puniceus Nutt.

Lyon (1886) CA.

Mimulus guttatus Fisch. ex DC. var. depauperatus (Gray)

Grant

Mimulus guttatus Fisch. ex DC. ssp. guttatus
Gentry (1949) based on Elmore 257 AHFH!, misiden-
tified, CR.

Mimulus latifolius Gray
Mimulus brandegei Penn.

Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890b) CR; Jepson (1925) CR; Munz (1935) CR;
Eastwood (1941) CR; Dunkle (1950) CR.

Mimulus longiflorus (Nutt.) Grant

Diplacus longiflorus Nutt. ssp. longiflorus
Munz (1935) CR; Dunkle (1950) RO, CR.

Mimulus longiflorus (Nutt.) Grant var. linearis (Benth.) Grant
Diplacus longiflorus Nutt. ssp. longiflorus
Dunkle (1950) CR.

Diplacus puniceus Nutt.

Dunkle (1950) CA.

Mimulus luteus L.

Mimulus guttatus Fisch. ex DC. ssp. guttatus
Lyon (1 886) CA; Brandegee ( 1888) RO, CR; Yates (1889)
CR; Brandegee (1890b) RO, CR, CA; Trask (1899)
CA.

Mimulus moschatus Dougl.
incertae sedis
Eastwood (1941) RO, CR.

Mimulus nasutus Greene

probably Mimulus guttatus Fisch. ex DC. ssp. guttatus
Greene (1887a) CR; Brandegee (1888) RO; Yates (1889)
RO, CR.

Mimulus puniceus (Nutt.) Steud.

Diplacus puniceus Nutt.

Davidson (1896) CA; Abrams and Ferris (1923-1960)
CA; Munz (1935) CA; Dunkle (1950) CA; Munz
and Keck (1959) CA; Thome (1967) CA; Munz
(1974) CA; Wiggins (1980) CA.

Mirabilis californica Gray

Mirabilis heimerlii (Standi.) Macbr.

Greene (1885) GU.

Mirabilis laevis (Benth.) Curran
Mirabilis californica Gray var. californica

Hoffmann (1932a) RO; Munz (1935) CR, CA; Dunkle
(1942) AN, BA; Dunkle (1950) RO, CR, BA, CA;
Munz and Keck (1959) CR, CA; Raven (1963) CL;
Thome (1967) CA; Philbnck (1972) BA.

Mirabilis heimerlii (Standi.) Macbr.

Franceschi (1893) GU; Moran (1951) GU.

Mirabilis laevis (Benth.) Curran var. cedrosensis (Standi.)
Munz

Mirabilis californica Gray var. californica

Munz (1935) CL; Dunkle (1950) CL; Munz and Keck
(1959) CL.

Mirabilis laevis (Benth.) Curran var. cordifolia Dunkle
Mirabilis californica Gray var. californica

Dunkle (194 1 ) based on Dunkle 7234 LAM! CL; Dunkle
(1950) CL; Munz and Keck (1959) CL.

Monardella lanceolata Gray
incertae sedis

Lyon (1886) CA; Brandegee (1890b) CA.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 119

Montia perfoliata (Donn) Howell

Claytonia perfoliata Donn var. perfoliata
Millspaugh and Nuttall ( 1 923) CA; Eastwood ( 1 929) GU;
Hoffmann (1932a) MI; Munz (1935) the islands;
Eastwood (1941) MI, RO, CR, CA; Dunkle (1942)
AN, BA; Howell (1942) GU; Gentry (1949) CL;
Dunkle (1950) MI, RO, CR, AN, NI, BA, CA, CL,
GU; Raven (1963) CL; Smith (1976) MI, RO, CR,
AN; Wiggins (1980) GU.

Montia perfoliata (Donn) Howell var. parviflora (Dougl. ex
Hook.) Jeps.

Claytonia perfoliata Donn var. parviflora (Dougl. ex Hook.)

Torr.

Hoffmann (1932a) CR; Eastwood (1941) CR; Smith
(1976) CR.

Muhlenbergia debilis (HBK.) Kunth
Muhlenbergia microsperma (DC.) Kunth

Watson (1876) GU; Greene (1885) GU; Greene (1887a)
CR; Yates ( 1 889) CR; Brandegee ( 1 890b) CR; Vasey
and Rose (1890) based on Palmer 656 ND-G! and
Palmer 670 ND-G! GU; Franceschi (1893) GU.

Muhlenbergia gracilis (HBK.) Kunth
Muhlenbergia microsperma (DC.) Kunth

Brandegee (1890a) CA; Brandegee (1890b) CA; David-
son (1896) CA.

Munzothamnus blairii (M. & J.) Raven
Stephanomeria blairii M. & J.

Raven (1963) CL; Thome (1969) CL; Munz (1974) CL.

Myosurus lepturas (Gray) Howell var. filiformis (Greene)
Abrams

Myosurus minimus L. var. filiformis Greene
Abrams and Ferris (1923-1960) GU.

Nasturtium aquaticum Tragus
Nasturtium officinale R. Br.

Greene (1887a) CR; Yates (1889) CR.

Navarretia filifolium (Nutt.) Kuntze
Eriastrum filifolium (Nutt.) Woot. & Standi.

Millspaugh and Nuttall (1923) CA; Eastwood (1941) CR,
CA.

Navarretia viscidula Benth.

Navarretia atractyloides (Benth.) H. & A.

Millspaugh and Nuttall (1923) CA; Eastwood (1941 )CA.

Nemophila aurita Lindl.

Pholistoma racemosum (Nutt.) Const.

Watson (1876) GU.

Nemophila erodiifolia Millsp. in Millsp. & Nutt.

Pholistoma racemosum (Nutt.) Const.

Millspaugh and Nuttall (1923) CA; Eastwood (1941) CA.

Nemophila parviflora Dougl. ex Benth.

probably Nemophila pedunculata Dougl. ex Benth.
Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890b) CR; Eastwood (1941) CR.

Nemophila pedunculata Dougl. ex Benth.

Hoffmann (1932b) based on Hoffmann s.n. Apr. 10,
1930 SBM#9226!, misidentified, MI, based on
Hoffmann s.n. Apr. 17, 1929 SBM#5992!,

SBM#5993! and SBM#5994!, misidentified, RO;
Eastwood (1941) MI, RO.

Nemophila racemosa Nutt, ex Gray
omitted as unsubstantiated on NI; also omitted by Fore-
man (1967).

Dunkle (1950) NI.

Neomammillaria goodridgei (Scheer) Britt. & Rose
Mammillaria blossfeldiana Boedecker var. shurliana Gates
Eastwood (1929) GU.

Nesothamnus incanus (Gray) Rydb.

Perityle incana Gray
Howell (1942) GU.

Nicotiana bigelovii (Torr.) Wats.

Nicotiana attenuata Torr. ex Wats, in King

Watson (1876) based on Palmer 64 NY!, misidentified,
GU.

Nicotiana clevelandii Gray
incertae sedis

Brandegee (1890a) CA; Eastwood (1941) CA.

Nicotiana petuniaeflora Greene
Nicotiana attenuata Torr. ex Wats, in King
Greene (1885) based on Greene s.n. Apr. 25, 1885
CAS#859! GU; Eastwood (1929) GU.

Notholaena californica D.C. Eat.
omitted as unsubstantiated on RO.

Eastwood (1941) RO.

Aspidotis californica (Hook.) Nutt, ex Copel.

Eastwood (1941) CR, CA.

Notholaena Candida (Mart. & Gal.) Hook.

Notholaena californica (D.C. Eat.) (A. Smith pers. comm.).
Greene (1887a) CR; Yates (1889) CR; Yates (1890) CR;
Dunkle (1940a) CR.

Notholaena newberryi D.C. Eat.
omitted as unsubstantiated on CR.

Eastwood (1941) CR.

Cheilanthes newberryi (D.C. Eat.) Domin (A. Smith pers.
comm.).

Watson (1876) GU; Greene (1885) GU; Lyon (1886)
CL; Brandegee (1890b) CL, GU; Vasey and Rose
(1890) GU; Franceschi (1893) GU; Abrams and
Ferris (1923-1960) CL, GU; Eastwood (1929) GU;
Dunkle (1940a) CL; Eastwood (1941) CL; Howell
(1942) GU; Dunkle (1950) CL, GU; Wiggins (1980)
GU.

Oenothera biennis L. var. hirsutissirna Gray ex Wats.
Oenothera elata HBK. ssp. hirsutissirna (Gray ex Wats.)
Dietrich

Brandegee (1890b) CR.

Oenothera bistorta Nutt, in T. & G.
incertae sedis

Lyon (1886) CA; Greene (1887a) CR; Greene (1887b)
MI; Brandegee (1888) RO; Yates (1889) MI, RO,
CR; Brandegee ( 1 890b) MI, RO, CR, CA; Eastwood
(1941) MI, RO, CR, CA.

Oenothera cheiranthifolia Homem. ex Spreng.

120 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

omitted as unsubstantiated on CA.

Eastwood (1941) CA; Gentry (1949) CA.

Camissonia cheiranthifolia (Homem. ex Spreng.) Raim. in
Engl. & Prantl ssp. cheiranthifolia
Greene (1887a) CR; Greene (1887b) MI; Brandegee
(1888) RO; Yates (1889) MI, RO, CR; Brandegee
( 1 890b) MI, RO, CR; Hoffmann (1932b) RO; Munz
(1935) MI, RO, BA; Eastwood (1941) MI, RO, CR,
NI; Dunkle ( 1 942) BA; Gentry ( 1 949) MI, RO, CR,
NI, BA; Dunkle (1950) MI, RO, CR, NI, BA; Munz
and Keck (1959) MI, RO; Raven (1963) CL.

Oenothera cheiranthifolia Homem. ex Spreng. var. nitida
(Greene) Munz

Camissonia cheiranthifolia (Homem. ex Spreng.) Raim. in
Engl. & Prantl ssp. cheiranthifolia
Jepson (1909-1943) MI; Abrams and Ferris (1923-1960)
MI; Hoffmann (1932b) RO; Munz (1935) MI, RO;
Dunkle (1950) RO, CR; Munz and Keck (1959) MI.

Oenothera contorta Dougl. in Hook. var. epilobioides (Greene)
Munz

Camissonia strigulosa (F. & M.) Raven

Hoffmann (1932b) MI; Eastwood (1941) MI, RO.

Oenothera contorta Dougl. in Hook. var. strigulosa (F. & M.)
Munz

Camissonia strigulosa (F. & M.) Raven
Hoffmann ( 1932b) RO; Munz (1935) RO; Dunkle ( 1950)
RO; Munz and Keck (1959) RO.

Oenothera dentata Cav.

Camissonia strigulosa (F. & M.) Raven

Brandegee (1888) RO; Yates (1889) RO; Brandegee
(1890b) RO.

Oenothera guadalupensis Wats.

Camissonia guadalupensis (Wats.) Raven ssp. Clementina
(Raven) Raven

Abrams and Ferris (1923-1960) CL; Munz (1935) CL;
Eastwood (1941) CL; Dunkle (1950) CL; Munz and
Keck (1959) CL.

Camissonia guadalupensis (Wats.) Raven ssp. guadalu-
pensis

Watson (1876) GU; Abrams and Ferris (1923-1960)
GU; Eastwood (1929) GU; Dunkle (1950) GU;
Munz and Keck (1959) GU.

Oenothera guadalupensis Wats. ssp. Clementina Raven

Camissonia guadalupensis (Wats.) Raven ssp. Clementina
(Raven) Raven
Raven (1963) CL.

Oenothera hirta Link. var. jonesii H. Lev.

Camissonia micrantha (Homem. ex Spreng.) Raven
Clokey (1931) CR.

Oenothera hookeri T. & G.

omitted as unsubstantiated on RO and CA.

Eastwood (1941) RO, CA.

Oenothera elata HBK. ssp. hirsutissima (Gray ex Wats.)
Dietrich

Greene (1887a) CR; Yates (1889) CR; Eastwood (1941)
CR.

Oenothera leptocarpa Greene

Camissonia californica (Nutt, ex T. & G.) Raven
Hoffmann (1932b) RO; Thorne (1967) CA.

Oenothera micrantha Homem. ex Spreng.

Camissonia micrantha (Homem. ex Spreng.) Raven
Lyon (1886) CA; Brandegee (1890b) CA; Trask (1899)
CA; Hoffmann (1932b) MI, RO, CR; Eastwood
( 1 94 1 ) MI, RO, CR, CA, CL; Munz and Keck (1959)
Channel Islands; Raven (1963) CL; Thome (1967)
CA.

Camissonia robusta Raven

Moran (1951) based on Moran 2891 RSA! cited by Ra-
ven (1969) GU.

Oenothera micrantha Homem. ex Spreng. var. jonesii (H.
Lev.) Munz

Camissonia hirtella (Greene) Raven
Munz and Keck (1959) Channel Islands.

Oenothera nitida Greene

Camissonia cheiranthifolia (Homem. ex Spreng.) Raim. in
Engl. & Prantl ssp. cheiranthifolia
Greene (1887b) MI; Yates (1889) MI; Eastwood (1941)
MI, RO.

Oenothera spiralis Hook. var. nitida (Greene) Jeps.

Camissonia cheiranthifolia (Homem. ex Spreng.) Raim. in
Engl. & Prantl ssp. cheiranthifolia
Jepson (1925) MI.

Oenothera strigulosa (F. & M.) T. & G.

Camissonia strigidosa (F. & M.) Raven
Eastwood (1941) RO.

Oenothera viridescens Hook.

Camissonia cheiranthifolia (Homem. ex Spreng.) Raim. in
Engl. & Prantl ssp. suffruticosa (Wats.) Raven
Eastwood (1898) NI.

Oligomeris glaucescens Camb.

Oligomeris linifolia (Vah.) Macbr.

Eastwood ( 1 898) NI; Millspaugh and Nuttall (1923) CA;
Eastwood (1941) MI, RO, CR, NI, CA, CL.
Opuntia chotla Weber
Opuntia prolifera Engelm.

Wiggins (1980) AN.

Opuntia engelmannii Salm-Dyck. in Engelm.

Opuntia littoralis (Engelm.) Ckll. var. littoralis
Lyon (1886) CA.

Opuntia engelmannii Salm-Dyck. in Engelm. var. littoralis
Engelm.

Opuntia littoralis (Engelm.) Ckll. var. littoralis
Greene (1887a) CR; Greene (1887b) MI; Brandegee
(1888) RO; Yates (1889) MI, RO, CR, AN; Bran-
degee (1890b) MI, RO, CR, CA; Eastwood (1898)
NI; Trask (1899) CA; Trask (1904) CL.

Opuntia megacantha Salm-Dyck.

Opuntia ficus-indica (L.) Mill.

Millspaugh and Nuttall ( 1 923) based on Millspaugh 4523
F!, misidentified CA; Jepson (1909-1943) CA.
Opuntia megacarpa Griff,
probably Opuntia ficus-indica (L.) Mill.

Eastwood (1941) CA.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 121

Opuntia occidentalis Engelm. & Bigel.

Opuntia littoralis (Engelm.) Ckll. var. littoralis

Millspaugh and Nuttall (1923) CA; Davidson and Mox-
ley (1923) CA; Howell (1935) NI; Eastwood (1941)
Ml, RO, CR, NI, CA, CL.

Opuntia phaecantha Engelm. in Gray var. discata (Griffiths)
Benson & Walkington
incertae sedis
Thome (1967) CA.

Orobanche californica Cham. & Schlecht.
probably Orobanche parishii (Jeps.) Heckard ssp. brachy-
loba Heckard.

Thome (1967) CA.

Orobanche fasciculata Nutt. var. franciscana Achey
Orobanche fasciculata Nutt.

Thome (1967) CA; Smith (1976) MI, RO.

Orobanche grayana G. Beck

possibly Orobanche californica Cham. & Schlecht. ssp.
grandis Heckard

Hoffmann (1932b) RO; Eastwood (1941) RO.
Orobanche uniflora L. var. minuta (Suksd.) Achey
Orobanche uniflora L. ssp. occidentalis (Greene) Abrams
ex Ferris

Munz and Keck (1959) CR.

Orobanche uniflora L. var. sedi (Suksd.) Achey
Orobanche fasciculata Nutt.

Munz (1935) based on Hoffmann s.n. May 8, 1932 POM!,
misidentihed, RO.

Orobanche uniflora L. ssp. occidentalis (Greene) Abrams
ex Ferris

Eastwood (1941) CR.

Orthocarpus purpurascens Benth.
incertae sedis

Brandegee (1890a) CA; Brandegee (1890b) CA; Trask
( 1 904) CA; Gentry ( 1 949) MI, CR in part, CA; Dun-
kle (1950) MI, CR, CA.

Orthocarpus densijlorus Benth. var. densiflorus
Eastwood ( 1 898) based on Trask 6 1 MO!, misidentihed,
NI; Eastwood (1941) NI; Gentry (1949) based on
Elmore 460 AHFH!, misidentihed, CR; Dunkle
(1950) NI.

Orthocarpus purpurascens Benth. var. pallidus Keck
Brandegee (1888) RO; Yates (1889) RO; Brandegee
( 1 890b) RO; Gentry ( 1 949) RO; Dunkle (1950) RO.
Oxalis californica (Abrams) Knuth
Oxalis albicans HBK. ssp. californicus (Abrams) Eiten
Abrams and Ferris (1923-1960) CA; Munz and Keck
(1959) CR, CA.

Oxalis pilosa Nutt.

Oxalis albicans HBK. ssp. pilosa (Nutt.) Eiten
Munz (1935) CR; Eastwood (1941) CR; Munz and Keck
(1959) CR.

Oxalis wrightii Gray

Oxalis albicans HBK. ssp. pilosa (Nutt.) Eiten
Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890b) CR.

Parietaria debilis Forst. f.

Parietaria hespera Hinton

Watson (1876) GU; Greene (1885) GU; Lyon (1886)
CA, CL; Greene (1887a) CR; Brandegee (1888) RO;
Yates (1889) RO, CR; Vasey and Rose (1890) GU;
Brandegee ( 1 890b) RO, CR, CA, CL; Jepson ( 1 909-
1943) CR; Millspaugh and Nuttall (1923) CA.

Parietaria floridana Nutt.

Parietaria hespera Hinton

Eastwood (1929) GU; Howell (1935) NI; Munz (1935)
RO, CR, CL; Eastwood (1941) RO, CR, NI, CA,
CL; Dunkle (1942) AN; Howell (1942) GU; Raven
(1963) CL; Thome (1967) CA.

Soleirolia soleirolii (Req.) Dandy
Foreman (1967) based on Foreman 120 UC!, misiden-
tihed, NI.

Parietaria hespera Hinton var. californica Hinton
Parietaria hespera Hinton
Smith (1976) RO, CR, AN.

Pelargonium sp.

Pelargonium peltatum (L.) L’Her. ex Ait.

Foreman (1967) based on Foreman 106 UC! NI.

Pe/laea mucronata (D.C. Eat.) D.C. Eat.
omitted as column transposition for CA.

Dunkle (1950) BA.

omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Dunkle (1950) CL.

Penstemon cordifolius Benth.

Keckiella cordifolia (Benth.) Straw
Lyon (1886) CA; Greene (1887a) CR; Brandegee (1888)
RO; Yates (1889) RO, CR; Ford (1890) CR; Bran-
degee ( 1 890b) RO, CR, CA; Trask ( 1 899) CA; Trask
(1904) CL; Millspaugh and Nuttall (1923) CA; Munz
(1935) RO, CR, CA, CL; Eastwood (1941) RO, CR,
CA, CL; Dunkle (1942) AN; Raven (1963) CL;
Thome (1967) CA; Smith (1976) RO, CR, AN.

Perityle emoryi Torr. in Emory
omitted as unsubstantiated on MI.

Dunkle (1950) MI.

Perityle fitchii Torr.

Perityle emoryi Torr. in Emory

Lyon (1886) CL; Greene (1887a) CR; Yates (1889) CR.

Perityle grayi Rose

Perityle emoryi Torr. in Emory

Vasey and Rose (1890) GU; Eastwood (1929) GU; How-
ell (1942) GU.

Petromecon frutescens Greene
Eschscholzia frutescens (Greene) J.T. Howell
Eastwood (1929) GU.

Petromecon palmeri (Rose) Greene
Eschscholzia palmeri Rose
Eastwood (1929) GU.

Peucedanum sp.
incertae sedis
Greene (1887a) CR.

122 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Peucedanum caruifolium (H. & A.) T. & G.

Lomatium caruifolium (H. & A.) Coult. & Rose

Brandegee (1888) RO, CR; Yates ( 1 889) RO, CR; Bran-
degee (1890b) RO, CR.

Phaca fastidia Kell.

Astragalus trichopodus (Nutt.) Gray ssp. leucopsis (T. &
G.) Thorne

Millspaugh and Nuttall (1923) CA.

Phaca leucopsis T. & G.

Astragalus trichopodus (Nutt.) Gray ssp. leucopsis (T. &
G.) Thome

Millspaugh & Nuttall (1923) CA.

Phacelia douglasii Torr.

Pholistoma racemosum (Nutt.) Const.

Yates (1889) CR; Jepson (1909-1943) CA.

Phacelia hispida Gray

omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Eastwood (1941) BA.

Phacelia insularis Munz

omitted as unsubstantiated on CR.

Dunkle (1950) CR.

Phacelia parryi Torr.

Phacelia viscida (Benth.) Torr.

Greene (1887a) CR; Yates (1889) CR; Eastwood (1941)
CR.

Phacelia phyllomanica Gray
Phacelia floribunda Greene

Lyon (1886) CL; Eastwood (1941) CL; Dunkle (1950)
CL.

Phacelia ramosissima Dougl. ex Lehm.
incertae sedis

Brandegee (1888) RO; Yates (1889) RO; Brandegee
(1890b) RO, CR.

Phacelia ramosissima Dougl. ex Lehm. var. cinerea (Eastw.
ex Macbr.) Jeps.

Phacelia cinerea Eastw. ex Macbr.

Jepson (1925) NI.

Phacelia suffrutescens Parry

Phacelia ramosissima Dougl. ex Lehm. var. austrolittoralis
Munz

Greene (1887a) based on Greene s.n. in Jul.-Aug. 1886
ND-G#042054!, misidentified, CR; Yates (1889)
CR; Eastwood (1941) CR.

Phacelia tanacetifolia Benth.

Phacelia cicutaria Greene ssp. bispida (Gray) J. Beau-
champ ex Thome

Munz (1974) probably based on Fosberg 7538 POM!,
misidentified, CR; Smith (1976) CR.

Phalaris bulbosa L.

Phalaris minor Retz.

Hoffmann (1932a) based on Hoffmann s.n. Jun. 15, 1930
SBM#7556!, misidentified, CR; Eastwood (1941)
CR.

Phalaris canariensis L.

omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Lyon (1886) CL.

Phalaris caroliniana Walt.

Eastwood (1898) based on Trask 9 MO! (annotated by
Anderson, 1959), misidentified, NI.

Phalaris intermedia Bose, ex Poir. in Lam.
possibly Phalaris caroliniana Walt.

Brandegee (1900) GU.

Phalaris caroliniana Walt.

Brandegee ( 1 888) based on Brandegee s.n. in Jun., 1 888
UC!, misidentified, CR; Yates (1889) CR; Bran-
degee (1890b) CR; Eastwood (1941) CR.

Phalaris minor Retz.

Phalaris caroliniana Walt.

Foreman (1967) based in part on Trask 9 MO! annotated
by Anderson (1959), misidentified, NI.

Philibertia hirtella (Gray) Parish
Sarcostemma cynanchoides Dene. ssp. hartwegii (Vail) R.
Holm.

Millspaugh and Nuttall ( 1 923) CA; Eastwood (1941) CA.
Phoradendron bolleanum (Seem.) Eichler
Phoradendron bolleanum (Seem.) Eichler ssp. densum
(Torr.) Wiens

Watson (1876) based on Palmer 85 CM! GU.

Phoradendron guadalupensis Trelease

Phoradendron bolleanum (Seem.) Eichler ssp. densum
(Torr.) Wiens
Eastwood (1929) GU.

Photinia arbutifolia (Ait.) Lindl. var. macrocarpa Munz
Heteromeles arbutifolia (Ait.) R. Roem.

Eastwood (1941) CA; Gentry (1949) CA, CL; Dunkle
(1950) CR, CA, CL; Munz and Keck (1959) CA,
CL.

Pinus insignis Dougl. ex Loud.

Pinus radiata D. Don var. binata (Engelm. in Wats.) Lem-
mon

Watson (1876) based on Palmer 90 NY! GU.

Pinus insignis Dougl. ex Loud. var. binata Engelm. in Wats.
Pinus radiata D. Don var. binata (Engelm. in Wats.) Lem-
mon

Greene (1885) GU; Brandegee (1890b) GU; Franceschi
(1893) GU.

Pinus remorata Mason

Greene (1887a) CR; Brandegee ( 1 888) RO; Yates ( 1 889)
RO, CR; Brandegee (1890b) RO, CR; Abrams and
Ferris (1923-1960) RO, CR.

Pinus radiata D. Don
incertae sedis

Jepson (1925) RO, CR; Gentry (1949) CR.

Pinus radiata D. Don var. binata (Engelm. in Wats.) Lem-
mon

Jepson (1925) GU; Howell (1942) GU.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 123

Pinus radiata D. Don f. binata (Engelm. in Wats.) J.T. Howell
Pinus radiata D. Don var. binata (Engelm. in Wats.) Lem-
mon

Howell (1942) based in part on Howell 8183 NY! GU.
Pinus radiata D. Don f. guadalupensis J.T. Howell

Pinus radiata D. Don var. binata (Engelm. in Wats.) Lem-
mon

Howell (1942) based in part on Howell 8267 CAS! GU.
Pinus radiata D. Don var. binata (Engelm. in Wats.) Lem-
mon

Pinus remorata Mason

Abrams and Ferris (1923-1960) RO; Wiggins (1980)
RO.

Pinus remorata Mason

Pinus radiata D. Don var. binata (Engelm. in Wats.) Lem-
mon

Wiggins (1980) GU.

Pityrogramma triangularis (Kaulf.) Maxon

omitted as column transposition for CA; also omitted by
Philbrick (1972).

Dunkle (1950, p. 293) BA.

Pityrogramma triangidaris (Kaulf.) Maxon var. semipallida
J.T. Howell
incertae sedis
Smith (1976) RO, CR.

Pityrogramma triangularis (Kaulf.) Maxon var. viscosa (D.C.
Eat.) Weath.

probably Pityrogramma triangidaris (Kaulf.) Maxon.
Dunkle (1950) GU; Wiggins (1980) GU.

Pityrogramma viscosa (D.C. Eat.) Maxon

omitted as column transposition for CA; also omitted by
Philbrick (1972).

Dunkle (1950, p. 293) BA.

Pityrogramma triangularis (Kaulf.) Maxon var. viscosa
(D.C. Eat.) Weath.

Millspaugh and Nuttall (1923) CA; Eastwood ( 1 94 1 ) RO,
CR, CA, CL; Dunkle (1950) RO, CR, CL.
Plagiobothrys arizonicus (Gray) Greene ex Gray
Plagiobothrys canescens Benth.

Brandegee (1890b) CA; Davidson (1896) CA; Mills-
paugh and Nuttall (1923) CA.

Plagiobothrys arizonicus (Gray) Greene ex Gray var. cata-
linensis Gray

Plagiobothrys canescens Benth.

Lyon (1886) CA; Davidson (1896) CA; Jepson ( 1 909—
1943) CA, CL; Davidson and Moxley (1923) CA;
Millspaugh and Nuttall (1923) CA; Eastwood (1941)
CA; Munz and Keck (1959) CA, CL.

Plagiobothrys californicus (Gray) Greene
incertae sedis

Greene (1887a) CR; Yates (1889) CR.

Plagiobothrys californicus (Gray) Greene var. californicus
Plagiobothrys californicus (Gray) Greene var. fulvescens

Jtn.

Munz and Keck (1959) probably based on Munz & Crow
1 1705 POM!, misidentified, RO; Smith (1976) RO.

Plagiobothrys californicus (Gray) Greene var. gracilis Jtn.
Plagiobothrys californicus (Gray) Greene var. californicus
Raven (1963) GU.

Plagiobothrys canescens Benth. var. catalinensis (Gray) Jeps.
Plagiobothrys canescens Benth.

Jepson (1925) CA.

Plagiobothrys cooperi Gray

Plagiobothrys californicus (Gray) Greene var. gracilis Jtn.
Brandegee (1890b) CA; Millspaugh and Nuttall (1923)
CA; Eastwood (1941) CR, CA.

Plagiobothrys tenellus (Nutt.) Gray
omitted as unsubstantiated on RO and erroneously attrib-
uted to Munz and Keck (1959).

Smith (1976) RO.

Plantago bigelovii Gray
Plantago erecta Morris ssp. erecta
Millspaugh and Nuttall (1923) based in part on Mill-
spaugh 4910 F!, and Nuttall 1215 F!, misidentified,
CA; Eastwood (1941) CA.

Plantago coronopus L. ssp. commutata (Guss.) Pilg.
Plantago coronopus L.

Abrams and Ferris (1923-1960) CA.

Plantago dura Morris
Plantago erecta Morris ssp. erecta
Davidson and Moxley (1923) CA; Millspaugh and Nut-
tall (1923) CA; Eastwood (1941) CA.

Plantago fastigiata Morris
Plantago ovata Forsk.

Thome (1969) CL.

Plantago heterophylla Nutt.

Plantago bigelovii Gray ssp. californica (Greene) Bassett
Munz (1935) MI, RO; Eastwood (1941) MI, RO.
Plantago hookeriana F. & M. var. californica (Greene) Poe.
Plantago erecta Morris ssp. erecta

Munz (1935) RO, CA, CL; Dunkle (1942) AN; Munz
and Keck (1959) Santa Barbara Islands.

Plantago insularis Eastw.

Plantago ovata Forsk.

Eastwood (1898) NI; Trask (1904) CL; Millspaugh and
Nuttall (1923) CA; Abrams and Ferris ( 1 923-1960)
Islands; Eastwood (1941) NI, CA, CL; Dunkle ( 1 942)
AN, BA; Howell ( 1 942) GU; Dunkle (1950) NI, BA,
CA, CL; Munz and Keck (1959) Santa Barbara Is-
lands; Raven (1963) CL; Foreman (1967) NI; Thome
(1967) CA; Munz (1974) Channel Islands; Smith
(1976) RO, CR “?,” AN.

Plantago insularis Eastw. var. insularis
Plantago ovata Forsk.

Wiggins (1980) GU.

Plantago maritima L.

omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Dunkle (1942) BA.

Plantago coronopus L.

Brandegee (1890a) CA.

124 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Plantago maritima L. var. California z (Fern.) Pilg.

Hoffmann (1932b) RO; Munz (1935) RO; Eastwood
(1941) RO.

Plantago obversa Morris
Plantago erecta Morris ssp. erecta
Davidson and Moxley (1923) CA.

Plantago patagonica Jacq.

Plantago erecta Morris ssp. erecta
Lyon (1886) CA; Greene (1887a) CR; Greene (1887b)
MI; Brandegee (1888) RO; Yates (1889) MI, RO,
CR; Brandegee (1890b) MI, RO, CR, CA.

Plantago ovata Forsk.

Watson (1876) based on Palmer 54 MO!, misidentified,
GU; Greene (1885) GU; Lyon (1886) CL; Vasey
and Rose (1890) based on Palmer 878 ND-G!, mis-
identified, GU; Brandegee (1890b) CL; Eastwood
(1929) GU.

Plantago speciosa Morris
Plantago erecta Morris ssp. erecta

Davidson and Moxley (1923) CA; Millspaugh and Nut-
tall (1923) CA; Eastwood (1941) AN, CA.
Platystemon aculeolatus Greene
Platystemon californicus Benth.

Jepson (1909-1943) based on type of Trask s.n. in May

1901 CAS#969! BA.

Platystemon californicus Benth.

omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Brandegee (1890b) CL.

Platystemon californicus Benth. var. ciliatus Dunkle
Platystemon californicus Benth.

Dunkle (1940b) based on Dunkle 7400 LAM! BA; Dun-
kle ( 1 942) BA; Dunkle (1950) BA; Munz (1974) BA.
Platystemon californicus Benth. var. nutans Bdg.
Platystemon californicus Benth.

Jepson (1909-1943) CR; Jepson (1925) CR; Dunkle
(1950) RO, CR; Munz (1974) RO, CR.
Platystemon californicus Benth. var. ornithopus (Greene)
Munz

Platystemon californicus Benth.

Dunkle (1950) MI, RO, CR, NI; Munz (1974) MI, RO,
NI.

Platystemon cernuus Greene
Platystemon californicus Benth.

Jepson (1909-1943) based on type of Trask s.n. on Mar.
1 897 CAS#966!, CA; Millspaugh and Nuttall ( 1 923)
CA; Eastwood (1941) CA.

Platystemon hispidulus Greene
Platystemon californicus Benth.

Jepson (1909-1943) based on type of Trask s.n. in Apr.
1897 CAS#958!, NI; Davidson and Moxley (1923)
NI, CA; Eastwood (1941) NI, CA.

Platystemon setosus Greene
Platystemon californicus Benth.

Jepson (1909-1943) based on type of Trask 1 1 in May

1902 CAS#971!, BA; Eastwood (1941) BA.

Platystigma californicum Benth. & Hook, in Brew. & Wats.
Meconella denticulata Greene
Brandegee (1890b) CR.

Platystigma denticulatum (Greene) Greene
Meconella denticulata Greene

Greene (1887a) CR; Yates (1889) CR.

Pluchea camphorata (L.) DC.

Pluchea odorata (L.) Cass.

Millspaugh and Nuttall (1923) CA; Abrams and Ferris
(1923-1960) CA; Hoffmann (1932b) CR; Munz
(1935) CR, CA; Eastwood ( 1941) CR, CA.

Pluchea purpurascens (Sw.) DC.

Pluchea odorata (L.) Cass. (Gillis, 1977).

Thome (1967) CA; Smith (1976) CR.

Poa nevadensis Vasey ex Scribn.
incertae sedis

Eastwood (1941) CR.

Poa steriantha Trin. (perhaps erroneous combination)
possibly Poa scabrella (Thurb.) Benth. ex Vasey.

Yates (1889) CR.

Polygonum aviculare L. var. littorale (Link) Koch
Polygonum aviculare L.

Hoffmann (1932a) CR; Eastwood (1941) RO, CR, CA.
Polygonum coccineum Muhl.

omitted as unsubstantiated on CA.

Thome (1967) CA.

Polygonum ramosissimum Michx.
incertae sedis

Hoffmann (1932a) CR; Eastwood (1941) CR.
Polypodium californicum Kaulf. var. kaulfussii D.C. Eat.
Polypodium californicum Kaulf.

Munz (1935) on islands; Dunkle (1940a) RO, CR, CA;
Eastwood (1941) RO, CR, CA, CL; Dunkle (1942)
AN, BA; Dunkle (1950) AN, BA, CA; Smith (1976)
AN.

Polypodium scouleri Hook. & Grev.

omitted as column transposition for CA; also omitted by
Philbrick (1972).

Dunkle (1950, p. 293) BA.

omitted as unsubstantiated on CA; also omitted by Mills-
paugh and Nuttall (1923).

Brandegee (1890a) CA; Brandegee (1890b) CA.
omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Dunkle (1950) CL.

Polypodium vulgare L. var. hesperium (Maxon) Nels. &
Macbr.

probably Polypodium californicum Kaulf.

Dunkle (1942) AN; Dunkle (1950) AN.

Polypodium vulgare L. var. kaulfussii (D.C. Eat.) Fern.
Polypodium californicum Kaulf.

Hoffmann (1932a) RO, CR.

Polystichum munitum (Kaulf.) Presl

Polystichum munitum (Kaulf.) Presl ssp. solitarium Maxon
(D.H. Wagner pers. comm.).

Watson (1876) based on Palmer 102 NY! GU; Eastwood

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 125

(1929) GU; Dunkle(1950) GU; Raven (1963) GU;
Smith (1976) GU; Wagner (1979) GU.

Populus fremontii Wats.

Populus x parryi Sarg.

Foreman (1967) based on Raven & Thompson 20728
DS!, misidentified, NI.

Populus fremontii Wats. var. wislizenii Wats.

Populus fremontii Wats. ssp. fremontii
Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890b) CR.

Potamogeton foliosus Raf.
omitted as unsubstantiated on CA.

Thome (1967) CA.

Potentilla anserina L.

Potentilla egedii Wormsk. var. grandis (Rydb.) J.T. Howell
Greene (1887b) MI; Yates (1889) MI; Eastwood (1941)
MI; Dunkle (1950) CR.

Primus ilicifolia (Nutt, ex H. & A.) Walp.

Prunus lyonii (Eastw.) Sarg.

Lyon (1886) CA; Brandegee (1890b) RO, CR, CA; Jep-
son (1925) Santa Barbara Islands; Eastwood (1941)
CA, CL; Dunkle (1950) RO, CR, CA, CL; Munz
and Keck (1959) CA, CL; Munz (1974) CA, CL;
Smith (1976) RO; Wiggins (1980) CA, CL.

Prunus ilicifolia (Nutt, ex H. & A.) Walp. ssp. lyonii (Eastw.)
Raven

Prunus lyonii (Eastw.) Sarg.

Raven (1963) CL; Thome ( 1 967) CA; Smith (1976) RO,
CR, AN.

Prunus ilicifolia (Nutt, ex H. & A.) var. occidentalis (Lyon)
Bdg.

Prunus lyonii (Eastw.) Sarg.

Brandegee (1888) RO, CR.

Prunus occidentalis Lyon
Prunus lyonii (Eastw.) Sarg.

Lyon (1886) CA; Greene (1887a) CR; Yates (1889) RO,
CR; Ford (1890) CR; Davidson (1896) CA.

Pteris aquilina L.

Pteridium aquilinum (L.) Kuhn var. pubcscens Underw.
Greene ( 1 887a) CR; Brandegee (1888) RO; Yates (1889)
RO, CR; Yates (1890) RO, CR; Brandegee (1890b)
RO, CR.

Quercus chrysolepis Liebm.

Quercus tomentella Engelm.

Watson (1876) GU.

Quercus dumosa Nutt. var. macdonaldii (Greene) Jeps.
Quercus x macdonaldii Greene
Jepson (1925) CR, CA.

Quercus dumosa Nutt. f. myrtifolia (Willd.) Trel.

Quercus dumosa Nutt.

Clokey (1931) CR.

Quercus lobata Nee
omitted as unsubstantiated on RO.

Brandegee (1888) RO.

Quercus x morehus Kell.

omitted as unsubstantiated on CA; also omitted by Mill-
spaugh and Nuttall (1923).

Davidson (1896) CA.

Quercus oblongifolia Torn.
incertae sedis

Brandegee (1890b) RO, CR, CA; Eastwood (1941) RO,
CR.

Ranunculus californicus Benth.
incertae sedis

Brandegee (1888) RO; Yates (1889) RO; Brandegee
(1890b) MI, RO, CR.

Ranunculus deppei Nutt, in T. & G.

Ranunculus californicus Benth. ssp. californicus
Greene (1887a) CR; Greene (1887b) MI; Yates (1889)
MI, CR.

Rhamnus catalinae A. Davids.

Rhamnus pirifolia Greene

Davidson and Moxley (1923) based on T. Payne 2344
LAM! CA; Eastwood (1941) CA.

Rhamnus crocea Nutt, in T. & G.

Rhamnus pirifolia Greene

Watson (1876) GU; Lyon (1 886) CA; Brandegee (1 890b)
MI, CR, CA; Trask (1899) CA; Trask (1904) CL;
Eastwood (1929) GU.

Rhamnus crocea Nutt, in T. & G. var. insularis (Greene)
Sarg.

omitted as unsubstantiated on BA; also omitted by Phil-
brick (1972).

Munz (1935) ambiguous implication perhaps a collec-
tive term for the Santa Barbara Islands, BA.
Rhamnus pirifolia Greene

Jepson (1909-1943) CR, CA; Hoffmann (1932b) RO;
Munz (1935) CA; Dunkle (1950) MI, RO, CR, CA,
CL.

Rhamnus crocea Nutt, in T. & G. ssp. pirifolia (Greene) C.B.
Wolf

Rhamnus pirifolia Greene

Abrams and Ferris (1923-1960) RO, CR, CA, CL, GU;
Munz and Keck (1959) Channel Islands.

Rhamnus insularis Greene
Rhamnus pirifolia Greene

Greene (1887a) CR; Greene (1887b) MI; Yates (1889)
MI, CR; Ford (1890) CR; Eastwood (1941) CR.
Rhamnus insulus Kell.

Rhamnus pirifolia Greene

Millspaugh and Nuttall (1923) CA; Eastwood (1941) MI,
RO, CR, CA, CL.

Rhus diversiloba T. & G.

Toxicodendron radicans (L.) Kuntze ssp. diversilobum (T.
& G.) Thome

Lyon (1886) CA; Greene (1887a) CR; Greene (1887b)
MI; Brandegee (1888) RO; Yates (1889) MI, RO,
CR; Brandegee (1890b) MI, RO, CR, CA; Trask
(1899) CA; Eastwood (1941) MI, RO, CR, CA;
Dunkle (1942) AN; Dunkle (1950) MI, RO, CR,
AN, CA, CL.

Rhus integrifolia (Nutt.) Benth. & Hook,
omitted as column transposition for CA; also omitted by
Philbrick (1972).

Dunkle (1950, p. 293) BA.

126 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Rhus laurina Nutt, in T. & G.

omitted as unsubstantiated on RO and CR.

Eastwood (1941) RO, CR; Dunkle (1950) RO, CR.
Malosma laurina (Nutt, in T. & G.) Nutt, ex Abrams
Watson (1876) GU; Greene (1885) GU; Lyon (1886)
CA; Brandegee ( 1 890b) CA; Franceschi (1893) GU;
Davidson (1896) CA; Trask (1899) CA; Jepson
( 1909-1 943) CA; Eastwood (1 929) GU; Munz(1935)
CA; Eastwood (1941) CA, CL; Dunkle (1950) CA,
CL; Munz (1974) CA.

Rhus ovata Wats.

omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Lyon (1886) CL.

Ribes malvaceum Sm. in Rees

omitted as unsubstantiated on CA.

Eastwood (1941) CA.

Ribes malvaceum Sm. in Rees var. clementinum Dunkle
Ribes malvaceum Sm. in Rees var. malvaceum

Dunkle (1943) based on Dunkle 7338 LAM! CL; Dunkle
(1950) CL.

Ribes menziesii Pursh
incertae sedis

Brandegee (1888) CR; Brandegee (1890b) CR; Munz
(1974) CR.

Ribes sanguineum Pursh

probably Ribes malvaceum Sm. in Rees var. malvaceum
Brandegee ( 1 890b) CR.

Ribes sanguineum Pursh var. malvaceum (Sm. in Rees) Loud.
Ribes malvaceum Sm. in Rees var. malvaceum

Brandegee (1888) CR; Yates (1889) CR; Ford (1890)
CR.

Ribes subvestitum H. & A.
omitted as unsubstantiated on CA.

Eastwood (1941) CA.

Ribes menziesii Pursh var. thacherianum Jeps.

Greene (1887a) CR; Yates (1889) CR; Ford (1890) CR;
Eastwood (1941) CR.

Ribes thacherianum (Jeps.) Munz
Ribes menziesii Pursh var. thacherianum Jeps.

Munz and Keck (1959) CR; Munz (1974) CR; Smith
(1976) CR.

Ribes viburniflorum Gray
Ribes viburnifolium Gray
Trask (1899) typographical error CA.

Rosa gratissima Greene
Rosa californica Cham. & Schlecht.

Gentry (1949) CR.

Rubus vitifolius Cham. & Schlecht.
omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Eastwood (1941) CL.

Rubus ursinus Cham. & Schlecht.

Millspaugh and Nuttall (1923) CA; Munz (1935) RO,
CR, CA; Eastwood (1941) MI, RO, CR; Dunkle
(1942) AN; Dunkle (1950) RO, CR, CA.

Rumex acetose/la L.

Rumex angiocarpus Murbeck

Hoffmann (1932a) based on Hoffmann s.n. Mar. 20,
1932 SBM#11767!, misidentified, CR; Eastwood
(1941) CR.

Rut a graveolens L.

Rut a chalepensis L.

Millspaugh and Nuttall (1923) CA; Eastwood (1941) CA.
Sagina apetala Ard.

Sagina decumbens (Ell.) T. & G. ssp. occidentalis (Wats.)
Crow

Crow (1978) based on Abrams & Wiggins 81
DS#209805!, misidentified (petals present, capsules
longer than sepals, leaf bases not ciliate), CR.
Sagina occidentalis Wats.

Sagina decumbens (Ell.) T. & G. ssp. occidentalis (Wats.)
Crow

Lyon ( 1 886) CA; Greene ( 1 887a) CR; Brandegee (1888)
RO; Yates ( 1889) RO, CR; Brandegee ( 1 890b) RO,
CR, CA; Millspaugh and Nuttall (1923) CA; Hoff-
mann (1932a) MI; Eastwood (1941) MI, RO, CR,
CA; Munz (1974) CR.

Salicornia sp.
incertae sedis
Greene (1887b) MI.

Salicornia bigelovii Torr.

probably Salicornia virginica L.

Eastwood (1941) MI.

Salicornia subterminalis Parish
Salicornia virginica L.

Dunkle (1942) based on Dunkle 7652 LAM!, misiden-
tified, AN; Gentry (1949) based on Elmore 402
AHFH!, misidentified, CL.

Salix argophylla Nutt.
incertae sedis

Millspaugh and Nuttall (1923) CA; Eastwood (1941) RO,
CR, CA.

Salix goodingii Ball
incertae sedis
Eastwood (1941) CR.

Salix laevigata Bebb.
incertae sedis
Brandegee (1888) RO.

Salix lasio/epis Benth. var. bigelovii (Torr.) Bebb.

Salix lasiolepis Benth.

Smith (1976) RO.

Salix lasiolepis Benth. var. bracelinae Ball
Salix lasiolepis Benth.

Foreman (1967) based on Raven & Thompson 20733
DS!, misidentified, NI.

Salix longifolia Muhl.
incertae sedis

Greene (1887a) CR; Yates (1889) CR; Ford (1890) CR;
Brandegee (1890b) CR.

Salsola kali L.

Salsola iberica Sennen & Pau
Foreman (1967) NI.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 127

Salsola pestifera A. Nels.

Salsola iberica Sennen & Pau
Thome (1967) CA.

Salvia apiana Jeps.
omitted as unsubstantiated on CR.

Eastwood (1941) CR.

Salvia brandegei Munz
omitted as unsubstantiated on AN.

Dunkle (1942) AN.

Salvia leucophylla Greene
incertae sedis
Eastwood (1941) CR.

Salvia mellifera Greene

omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Munz (1935) CL; Dunkle (1950) CL.

Salvia mellifera Greene var. jonesii Munz
omitted as unsubstantiated on MI and AN.

Eastwood (1941) (perhaps a misinterpretation of San
Miguel Mt. in San Diego Co. cited by Munz 1935)
MI; Dunkle (1942) AN; Dunkle (1950) CR.

Salvia palmeri (Gray) Greene
probably Salvia mellifera Greene.

Eastwood (1941) CR.

Sambucus caerulea Raf.

Sambucus mexicana Presl ex DC.

Millspaugh and Nuttall (1923) CA.

Sambucus glauca Nutt, in T. & G.

Sambucus mexicana Presl ex DC.

Lyon (1886) CA; Greene (1887a) CR; Brandegee (1888)
RO; Yates (1889) RO, CR; Brandegee (1890b) RO,
CR, CA; Trask (1899) CA; Trask (1904) CL.
Samolus floribundus HBK.

Samolus parviflorus Raf.

Jepson (1909-1943) CR; Abrams and Ferris (1923-1960)
CR.

Samolus valerandi L.

Samolus parviflorus Raf.

Brandegee (1890b) CR.

Samolus valerandi L. var. americanus Gray
Samolus parviflorus Raf.

Greene (1887a) CR; Yates (1889) CR; Eastwood (1941)
CR.

Sanicula bipinnatifida Dougl. ex Hook.

Sanicula arguta Greene ex Coult. & Rose

Lyon ( 1 886) CL; Brandegee ( 1 890b) CL; Davidson ( 1 896)
CL.

Sanicula laciniata H. & A.

Sanicula arguta Greene ex Coult. & Rose
Greene (1887a) CR; Yates ( 1 889) CR; Brandegee ( 1 890a)
CA; Brandegee (1890b) CR, CA; Millspaugh and
Nuttall (1923) CA.

Sanicula menziesii H. & A.

Sanicula arguta Greene ex Coult. & Rose
Eastwood (1898) NI.

Saxifraga malvaefolia Greene
Jepsonia malvaefolia (Greene) Small
Greene (1887a) CR; Yates (1889) CR.

Saxifraga parry i Torn
Jepsonia malvaefolia (Greene) Small
Brandegee (1890b) CR.

Saxifraga reflexa Hook.
incertae sedis

Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890b) CR.

Scandix pecten-veneris L.

Daucus pusillus Michx.

Foreman (1967) based on Foreman 230 UC!, misiden-
tified, NI.

Scirpus pungens Vahl.

Scirpus americanus Pers. var. monophyllus (Presl) Koya-
ma

Brandegee (1888) RO; Yates (1889) RO; Brandegee
(1890b) RO; Eastwood (1941) RO.

Scirpus riparius Spreng.
omitted as unsubstantiated on CR.

Yates (1889) CR; Eastwood (1941) CR.

Scirpus cernuus Vahl. ssp. californicus (Torn) Thome
Greene (1887b) MI; Yates (1889) CR; Brandegee (1890b)
MI; Eastwood (1941) MI, RO, CR.

Scrophularia californica Cham. & Schlecht.

Scrophularia villosa Penn, in Millsp. & Nutt.

Lyon (1886) CA; Brandegee (1890b) CA; Trask (1899)
CA.

Scrophularia californica Cham. & Schlecht. var. catalinae
Jeps.

omitted as unsubstantiated on MI and AN.

Dunkle (1942) AN; Dunkle (1950) MI, AN.
Scrophularia villosa Penn, in Millsp. & Nutt.

Munz (1935) CA, CL; Dunkle (1950) CA, CL.
Scutellaria tuberosa Benth. var. similis Jeps.

Scutellaria tuberosa Benth. ssp. australis Epl.

Smith (1976) CR.

Secale cereale L.

Elymus glaucus Buckl. ssp. glaucus

Thome (1967) based on Fosberg S4555 LAM!, misi-
dentified, CA.

Selaginella rupestris (L.) Spring.

Selaginella bigelovii Underw.

Lyon (1886) CA; Brandegee (1888) CR; Yates (1889)
CR; Brandegee (1890b) CR, CA.

Senecio lyonii Gray

omitted as unsubstantiated on CR, BA, and GU.
Eastwood (1941) CR; Gentry ( 1 949) CR, BA; also omit-
ted by Philbrick (1972); Dunkle (1950) GU.

Sida hederacea (Dougl.) Torr.

Malvella leprosa (Ortega) Krapovickas

Munz (1935) CL; Raven (1963) CL; Thome (1967) CA.
Sida leprosa (Ortega) K. Schum. var. hederacea (Dougl.) K.
Schum.

Malvella leprosa (Ortega) Krapovickas
Clement (1957) GU; Smith (1976) RO.

Sidalcea malvaeflora (DC.) Gray ex Benth.
omitted as unsubstantiated on CA.

Eastwood (1941) CA; Gentry (1949) CA.

128 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Silene conoidea L.

Silene multinervia Wats.

Brandegee (1888) CR; Yates (1889) CR; Brandegee
(1890a) CA.

Silene quinquevulnera L.

Silene multinervia Wats.

Greene (1887a) CR; Yates (1889) CR.

Sisymbrium canescens Nutt.

Descurainia pinnata (Walt.) Britt, ssp. menziesii (DC.) Detl.

Watson (1876) GU; Greene (1885) GU; Brandegee
(1890a) CA; Brandegee (1890b) CR, CA; Eastwood
(1929) GU.

Sisymbrium pinnatum (Walt.) Greene
Descurainea pinnata (Walt.) Britt, ssp. menziesii (DC.) Detl.

Greene (1887a) CR; Yates (1889) CR; Jepson (1909-
1943) CR; Hoffmann (1932b) RO.

Sisymbrium reflexum Nutt.

Caulanthus lasiophyllus (H. & A.) Pays.

Watson (1876) GU; Greene (1885) GU; Lyon (1886)
CL; Brandegee (1888) RO; Yates ( 1 889) RO; Vasey
and Rose (1890) GU; Brandegee (1890a) CA; Bran-
degee (1890b) MI, RO, CR, CA, CL.

Solanum calvum Bitter
Solanum nodiflorum Jacq.

Eastwood (1929) based in part on Palmer 60 in part NY!
GU.

Solanum nigrum L.

probably Solanum douglasii Dunal in DC.

Lyon (1886) CA; Brandegee (1890b) MI, RO, CR, CA.
possibly Solanum nodiflorum Jacq.

Franceschi (1893) GU.

Solanum nodiflorum Jacq.

Watson (1876) based on Palmer 60 in part NY! GU;
Vasey and Rose (1890) based on Palmer 860 NY!
GU.

Solanum nigrum L. var. douglasii (Dunal in DC.) Gray
Solanum douglasii Dunal in DC.

Watson (1876) based on Palmer 61 NY! GU; Brandegee
(1888) RO.

Solanum profundeincisum Bitter
Solanum douglasii Dunal in DC.

Eastwood (1929) based in part on Palmer 61 NY! GU.

Solanum villosum (L.) Mill.

Solanum douglasii Dunal in DC.

Gentry (1949) based on Elmore 200 AHFH! RO, based
on Elmore 420 AHFH! CL.

Solanum wallacei (Gray) Parish
Solanum wallacei (Gray) Parish ssp. clokeyi (Munz) Thome

Eastwood (1941) RO, CR.

Solanum wallacei (Gray) Parish ssp. wallacei

Wiggins (1980) CA, GU.

Solanum xantii Gray

Solanum wallacei (Gray) Parish ssp. clokeyi (Munz) Thome

Munz (1935) RO; Munz and Keck (1959) RO, CR; Smith
(1976) RO, CR.

Solanum wallacei (Gray) Parish ssp. wallacei

Watson (1876) based on Palmer 62 NY!, misidentified,
GU.

Solanum xantii Gray var. wallacei Gray

Solanum wallacei (Gray) Parish ssp. clokeyi (Munz) Thome
Greene (1887a) CR; Brandegee (1888) RO; Yates (1889)
RO, CR; Brandegee (1890b) RO, CR.

Solanum wallacei (Gray) Parish ssp. wallacei
Greene (1885) GU; Franceschi (1893) based on Fran-
ceschi 15 SBM! GU.

Sollya fusiformis Briq.

Sollya heterophylla Lindl.

Thome (1967) CA.

Sonchus tenerrimus L.

Sonchus asper (L.) Hill

Hoffmann (1932b) based on Hoffmann s.n. Mar. 25,
1932 SBM#1069!, misidentified, MI.

Sophia pinnata (Walt.) Howell
Descurainia pinnata (Walt.) Britt, ssp. menziesii (DC.) Detl.
Millspaugh and Nuttall (1923) CA.

Spergularia biflora (R. & P.) F. & M.
omitted as unsubstantiated on MI.

Dunkle (1950) MI.

Spergularia perfoliata (L.) DC.
incertae sedis

Greene (1887a) CR; Yates ( 1 889) CR; Brandegee ( 1 890b)
CR; Eastwood (1941) CR.

Spergularia marina (L.) Griseb.

omitted as column transposition for CA.

Dunkle (1950, p. 293) BA.

Spergidaria rubra (L.) J. & C. Presl

Spergidaria bocconei (Scheele) Foucaud
Eastwood (1941) CL.

Sphaeralcea fasciculata (Nutt.) Arthur var. laxiflora (Gray)
Jeps.

Malacothamnus fasciculatus (Nutt.) Greene ssp. catali-
nensis (Eastw.) Thome
Jepson (1909-1943) CA.

Sphaerostigma bistortum (Nutt, ex T. & G.) Walp.
Camissonia robusta Raven
Millspaugh and Nuttall (1923) CA.

Sphaerostigma micranthum (Homem. ex Spreng.) Walp.
Camissonia micrantha (Homem. ex Spreng.) Raven
Millspaugh and Nuttall (1923) CA.

Statice arctica Blake var. vulgaris Blake
Armeria maritima (Mill.) Willd. ssp. californica (Boiss.)
G.H.M. Lawr.

Munz (1935) RO; Dunkle (1950) RO.

Stellaria nitens Nutt.

omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Dunkle (1950) CL.

Stephanomeria elata Nutt.

Stephanomeria virgata Benth. ssp. virgata
Greene (1887a) CR; Yates (1889) CR.

Stephanomeria paniculata Nutt.

Stephanomeria virgata Benth. ssp. virgata

Lyon (1886) CA; Brandegee (1890b) CA; Eastwood
(1941) CA.

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 129

Stephanomeria tomentosa Greene
Stephanomeria virgata Benth. ssp. virgata (Gottlieb, 1972).
Greene (1887a) CR; Yates (1889) CR; Jepson (1925)
CR; Munz (1935) CR; Eastwood (1941) RO, CR;
Dunkle (1950) RO, CR.

Stephanomeria virgata Benth. var. tomentosa (Greene) Munz
Stephanomeria virgata Benth. ssp. virgata
Munz and Keck (1959) CR; Smith (1976) RO CR.
St ip a sp.
incertae sedis
Greene (1887a) CR.

Stipa andersonii (Vasey) Bdg.

Stipa lepida Hitchc.

Brandegee (1890b) RO, CR.

Stipa lepida Hitchc. var. andersonii (Vasey) Hitchc.

Stipa lepida Hitchc.

Millspaugh and Nuttall (1923) CA, Munz (1935) RO;
Eastwood (1941) RO, CA.

Stipa robusta (Vasey) Scribn.
incertae sedis
Eastwood (1898) NI.

Stipa vaseyi Scribn.

probably Stipa Columbiana Macoun var. nelsoni (Scribn.)
Hitchc.

Jepson (1909-1943) probably based on Trask 11 US!,
misidentified, NI; Davidson and Moxley (1923) NI;
Abrams and Ferris (1932-1960) NI.

Stipa viridula Trin.
incertae sedis

Brandegee (1888) CR; Yates (1889) CR.

Stylophyllum hassei Rose
Dudleya hassei (Rose) Moran
Davidson and Moxley (1923) CA; Millspaugh and Nut-
tall (1923) CA; Abrams (1917) CA; Abrams and
Ferris (1923-1960) CA; Eastwood (1941) CA.
Suaeda sp.

probably Suaeda californica Wats.

Eastwood (1898) NI.

Suaeda californica Wats. var. pubescens Jeps.

Suaeda californica Wats.

Dunkle (1950) MI, RO, CR, AN, NI, BA, CA, CL; Smith
(1976) RO, CR.

Suaeda californica Wats. var. taxifolia (Standi.) Munz
Suaeda californica Wats.

Smith (1976) RO, CR.

Suaeda fruticosa (L.) Forsk.

Suaeda californica Wats.

Howell (1942) based on Howell 8189 CAS!, misidenti-
fied, GU.

Suaeda taxifolia Standi.

Suaeda californica Wats.

Gentry (1949) based on Elmore 302 AHFH! BA.
Suaeda torreyana Wats.

probably Suaeda californica Wats.

Greene (1887a) CR; Greene (1887b) MI; Brandegee
(1888) RO; Yates (1889) MI, CR; Brandegee ( 1 890a)
CA; Brandegee ( 1 890b) MI, RO, CR, CA; Eastwood

(1941) MI, RO, CR, CA; Dunkle (1950) MI, RO,
CR, BA, CA.

Symphoricarpos albus (L.) Blake var. mollis (Nutt, in T. &
G.) Keck

Symphoricarpos mollis Nutt, in T. & G.

Munz (1935) CR, CA.

Syrmatium ornithopum (Greene) Greene
omitted as unsubstantiated on CR.

Eastwood (1941) CR.

Lotus argophyllus (Gray) Greene ssp. ornithopus (Greene)
Raven

Abrams (1917) NI, BA, CA, CL, GU; Millspaugh and
Nuttall (1923) CA; Eastwood (1941) NI, BA, CA,
CL.

Syrmatium patens Greene

Lotus scoparius (Nutt, in T. & G.) Ottley var. veatchii
(Greene) Ottley

Greene(1887a) CR; Greene(1887b) MI; Eastwood ( 1 94 1 )
MI.

Syrmatium traskiae Eastw. ex Noddin in Abrams
probably Lotus scoparius (Nutt, ex T. & G.) Ottley var.
dendroideus (Greene) Ottley
Davidson and Moxley (1923) CA; Millspaugh and Nut-
tall (1923) CA; Eastwood (1941) CA.

Lotus scoparium (Nutt, ex T. & G.) Ottley ssp. traskiae
(Eastw. ex Noddin in Abrams) Raven
Abrams (1917) CL; Davidson and Moxley (1923) CL;
Eastwood (1941) CL.

Syrmatium venustum (Eastw.) Davids. & Mox.

Lotus argophyllus (Gray) Greene ssp. ornithopus (Greene)
Raven

Davidson and Moxley (1923) NI.

Taraxacum officinale Wiggers
omitted as unsubstantiated on CL; also omitted by Raven
(1963).

Eastwood (1941) CL.

Taraxacum taraxacum (L.) Karst.

Taraxacum officinale Wiggers
Millspaugh and Nuttall (1923) CA.

Tellima affinis (Gray) Gray
Lithophragma affine Gray ssp. mixtum R.L. Taylor
Brandegee (1890a) CA; Brandegee (1890b) CA.
Tetragonia expansa Murr.

Tetragonia tetragonioides (Pall.) Kuntze
Hoffmann (1932a) MI, RO; Munz (1935) MI; Eastwood
(1941) MI, RO.

Thalesia fasciculata (Nutt.) Britt.

Orobanche fasciculata Nutt.

Millspaugh and Nuttall (1923) CA.

Thelesperma gracile (Torr.) Gray
Thelesperma megapotamicum (Spreng.) Kuntze
Millspaugh and Nuttall (1923) based in part on Nuttall
602 F! CA; Eastwood (1941) CA.

Thelypodium laciniatum (Hook.) Endl.
probably Caulanthus lasiophyllus (H. & A.) Pays.

Brandegee (1890a) CA; Brandegee (1890b) CA; Yates
(1889) CR.

130 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Thelypodium lasiophyllum (H. & A.) Greene

Caulanthus lasiophyllus (H. & A.) Pays.

Greene (1887a) CR; Greene (1887b) MI; Yates (1889)
MI, CR; Millspaugh and Nuttall (1923) CA; East-
wood ( 1 929) GU; Hoffmann ( 1 932b) MI, RO; East-
wood (1941) MI, RO, CR, AN, CA, CL; Thome
(1967) CA; Philbrick (1972) BA; Smith (1976) MI,
RO, CR, AN.

Thelypodium lasiophyllum (H. & A.) Greene var. inalienum

Rob.

Caulanthus lasiophyllus (H. & A.) Pays.

Smith (1976) AN.

Thysanocarpus conchuliferus Greene

omitted as unsubstantiated on CA.

Eastwood (1941) CA.

Thysanocarpus laciniatus Nutt, ex T. & G. var. ramosus
(Greene) Munz

omitted as unsubstantiated on CA.

Dunkle (1950) CA.

Tillaea angustifolia Nutt, in T. & G.

Crassula erect a (H. & A.) Berger
Brandegee (1890a) CA; Brandegee (1890b) CA; David-
son (1896) CA; Millspaugh and Nuttall (1923) CA.

Tillaea erect a H. & A.

Crassula erecta (H. & A.) Berger
Millspaugh and Nuttall (1923) CA; Eastwood (1929) GU;
Hoffmann (1932b) MI; Howell (1935) NI; Eastwood
(1941) MI, RO, CR, NI, CA; Dunkle (1942) BA;
Howell (1942) GU; Dunkle (1950) MI, RO, CR,
AN, NI, BA, CA, CL, GU.

Tillaea leptopetala Benth.

Crassula erecta (H. & A.) Berger
Vasey and Rose (1890) GU.

Tillaea minima Miers ex H. & A. in Hook.

Crassula erecta (H. & A.) Berger
Watson (1876) GU; Greene (1885) GU; Lyon (1886)
CA; Greene (1887a) CR; Brandegee (1888) RO;
Yates (1889) RO, CR; Brandegee (1890b) RO, CR,
CA.

Tissa macrotheca (Homem.) Britt.

Spergularia macrotheca (Homem.) Heynh. ssp. macro-
theca

Vasey and Rose (1890) GU; Brandegee (1890b) MI, RO,
CR, CA, CL; Millspaugh and Nuttall (1923) CA;
Eastwood (1929) GU.

Tissa pallida Greene ex Britt.

Spergularia macrotheca (Homem.) Heynh. ssp. macro-
theca

Vasey and Rose (1890) GU; Eastwood (1929) GU.

Tissa salina (J. & C. Presl) Britt.

Spergularia marina (L.) Griseb.

Brandegee (1890b) CA; Millspaugh and Nuttall (1923)
CA; Davidson and Moxley (1923) CA.

Tithymalus leptocerus (Engelm.) Millsp. in Millsp. & Nutt.

Euphorbia crenulata Engelm.

Millspaugh and Nuttall (1923) based on Millspaugh 4867
F!, Nuttall 170 F!, Nuttall 236 F! CA.

Toxicodendron diversilobum (T. & G.) Greene

Toxicondendron radicans (L.) Kuntze ssp. diversilobum (T.
& G.) Thome

Millspaugh and Nuttall (1923) CA; Gentry (1949) MI,
RO, CR, CA; Raven (1963) CL; Smith (1976) ap-
parently all four Channel Islands.

Trichosterigma miserum (Benth.) Kl. & Gke.

Euphorbia misera Benth.

Davidson and Moxley (1923) CA; Millspaugh and Nut-
tall (1923) CA.

Trifolium amplectens T. & G.
incertae sedis

Greene (1887a) CR; Brandegee (1890a) CA; Brandegee
( 1 890b) CR, CA; Davidson ( 1 896) CA; Trask ( 1 899)
CA; Jepson (1909-1943) CR, CA; Dunkle (1950)
MI, RO, CR, NI, CA, CL, GU.

Trifolium dichotomum H. & A.

Trifolium albopurpureum T. & G.

Eastwood (1898) based on Trask 37 LAM!, NY! NI;
Foreman (1967) based in part on Trask 37 LAM!,
NY! NI.

Trifolium fucatum Lindl. var. flavulum Jeps.

Trifolium fucatum Lindl. var. gambelii (Nutt.) Jeps.

Hoffmann (1932b) MI; Munz (1935) MI; Eastwood
(1941) MI, RO, CR.

Trifolium gracilentum T. & G.

Trifolium palmeri Wats.

Dunkle (1950) probably based on Dunkle 7415 LAM!
and Dunkle 7449 AHFH!, misidentified, BA.

Trifolium gracilentum T. & G. var. inconspicuum Fern.
Trifolium gracilentum T. & G.

Hoffmann (1932b) MI, RO; Eastwood (1941) MI, RO;
Dunkle (1950) RO, CR.

Trifolium gracilentum T. & G. var. palmeri McDer.
omitted as unsubstantiated on CR.

Dunkle (1950) CR.

Trifolium microdon H. & A.

Trifolium tridentatum Lindl. var. aciculare (Nutt.) McDer.

Dunkle (1942) probably based on Dunkle 7425 LAM!,
misidentified, BA; Dunkle (1950) BA.

Trifolium stenophyllum Nutt.
incertae sedis

Eastwood (1898) NI; Eastwood ( 1 94 1 ) NI; Dunkle (1950)
NI.

Trifolium stenophyllum Nutt.

Trifolium amplectens T. & G. var. truncatum (Greene)
Jeps.

Nuttall (1848) CA; Millspaugh and Nuttall (1923) CA;
Eastwood (1941) MI, RO, CR, CA, CL; Dunkle
(1950) MI, RO, CR, CA, CL.

Trifolium depauperatum Desv.

Howell (1935) based on Howell 8212 CAS!, misidenti-
fied, NI.

Trifolium tridentatum Lindl.
incertae sedis

Greene (1887a) CR; Greene (1887b) MI; Brandegee
(1888) RO; Yates (1889) CR; Brandegee (1890a)

Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands 131

CA; Brandegee (1890b) MI, RO; Trask (1899) CA;
Trask (1904) CL; Jepson (1909-1943) CR, CA.

Trisetum barbatum Steud.

Bromus trinii Desv.

Brandegee (1888) based on Brandegee s.n. in 1888 UC!
CR; Yates ( 1 889) CR; Brandegee ( 1 890b) CR; Vasey
and Rose (1890) based in part on Palmer 658 US!,
and Palmer 667 US!, UC! GU; Eastwood (1941)
CR.

Triteleia lugens Greene
Triteleia guadalupensis Lenz

Moran (1951) GU.

Tropidocarpum dubium A. Davids.

Tropidocarpum gracile Hook.

Eastwood (1941) CL.

Typha sp.
incertae sedis

Hoffmann (1932a) RO.

Typha angustifolia L.

Typha domingensis Pers.

Jepson (1909-1943) CR; Millspaugh and Nuttall (1923)
CA; Eastwood (1941) RO, CR, CA; Foreman (1967)
based on Blakley 4158 SBBG!, misidentified, NI.

Typha domingensis Pers.

Typha latifolia L.

Raven (1963) based on Raven 18018 RSA!, misidenti-
fied, CL.

Urtica gracilis Ait. var. holoserica (Nutt.) Jeps.

Urtica dioica L. ssp. holoserica (Nutt.) Thome

Munz (1935) CR, CA; Dunkle (1942) AN.

Urtica holoserica Nutt.

Urtica dioica L. ssp. holoserica (Nutt.) Thome

Lyon (1886) CA; Greene (1887a) CR; Yates (1889) CR;
Brandegee ( 1 890b) CR, CA; Millspaugh and Nuttall
( 1 923) CA; Eastwood ( 1 94 1 ) CR, CA; Gentry ( 1 949)
CR, CA; Munz and Keck (1959) CR, CA; Munz
(1974) CR, CA; Smith (1976) CR.

Uva-ursi pungens (HBK.) Abrams
Arctostaphylos catalinae P.V. Wells

Millspaugh and Nuttall (1923) CA.

Uva-ursi tomentosa (Pursh) Abrams
Arctostaphylos catalinae P.V. Wells

Millspaugh and Nuttall (1923) CA.

Verbena prostrata R. Br. in Ait.
incertae sedis

Lyon (1886) CA; Greene (1887a) CR; Greene (1887b)
MI; Brandegee (1890b) CA; Millspaugh and Nuttall
(1923) CA.

Verbena robusta Greene
Verbena lasiostachys Link

Jepson (1909-1943) CL; Eastwood (1941) CL; Gentry
(1949) CL.

Vicia americana Muhl. ex Willd. var. linearis Wats.
incertae sedis

Jepson (1909-1943) CA; Eastwood (1941) CA.

Vicia hassei Wats.

Jepson (1909-1943) based on Munz 6669 POM!, UC!
CL.

Vicia californicus Greene
Vicia americana Muhl. ex Willd.

Hoffmann (1932b) based on Hoffmann s.n. Apr. 16,
1929 SBM#6195! RO.

Vicia exigua Nutt, in T. & G.
omitted as unsubstantiated on NI; also omitted by Fore-
man (1967).

Dunkle (1950) NI.

Vicia exigua Nutt, in T. & G. var. hassei (Wats.) Jeps.

Vicia hassei Wats.

Wiggins (1980) GU.

Vinca minor L.

Sollya heterophylla Lindl.

Millspaugh and Nuttall (1923) based on Nuttall 801 F!,
misidentified, CA.

Vitis californica Benth.

Vitis girdiana Munson

Brandegee (1890a) CA; Brandegee (1890b) CA.

Vulpia megalura (Nutt.) Rydb.

Vulpia myuros (L.) K.C. Gmelin var. hirsuta Hack.
Philbrick (1972) BA.

Woodwardia radicans (L.) Sm.

Woodwardia fimbriata Sm. in Rees
Greene (1887a) CR; Yates (1889) CR; Yates (1890) CR;
Brandegee (1890b) CR; Hoffmann (1932a) RO.
Xanthium canadense Mill.

Xanthium strumarium L. var. canadense (Mill.) T. & G.
Greene (1887a) CR; Yates (1889) CR; Brandegee (1890b)
CR.

Xanthium pennsylvanicum Wallr.

omitted as unsubstantiated on MI and BA.

Eastwood (1941) MI, BA.

Xanthium strumarium L. var. canadense (Mill.) T. & G.
Millspaugh and Nuttall (1923) CA; Eastwood (1941) CR.
Xanthoxalis californica Abrams

Oxalis albicans HBK. ssp. californica (Abrams) Eiten
Millspaugh and Nuttall (1923) CA.

Zannichellia palustris L.

Ruppia maritima L.

Munz (1935) based on Wolf 3591 DS!, misidentified,
CA; Eastwood (1941) CA.

Zauschneria californica Presl
Epilobium canum (Greene) Raven ssp. canum

Lyon (1886) CA; Greene (1887a) CR; Greene (1887b)
MI; Brandegee (1888) RO; Brandegee (1890b) MI,
RO, CR, CA; Trask (1899) CA; Millspaugh and
Nuttall (1923) CA; Eastwood (1941) MI, CR, CA;
Dunkle (1950) RO, CA, CL; Munz and Keck (1959)
Santa Barbara Islands.

Zauschneria californica Presl ssp. angustifolia Keck
Epilobium canum (Greene) Raven ssp. canum
Abrams and Ferris (1923-1960) CA; Dunkle (1950) CA;
Munz and Keck (1959) CA.

Zauschneria californica Presl ssp. californica
Epilobium canum (Greene) Raven ssp. canum
Thome (1967) CA; Smith (1976) MI, RO, CR, AN.

132 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

Zauschneria californica Presl ssp. mexicana (Presl) Raven
Epilobium canum (Greene) Raven ssp. canum

Raven (1963) CL; Thorne (1967) CA; Smith (1976) CR.

Zauschneria californicum Presl var. villosa (Greene) Jeps.
Epilobium canum (Greene) Raven ssp. canum
Jepson ( 1 909- 1 943) RO, CR, CA, CL; Abrams and Fer-
ris ( 1 923-1960) RO, CR, CL; Munz ( 1 935) RO, CR,
CL; Gentry ( 1 949) RO, CR, CA, CL; Dunkle (1950)
RO, CR, CA, CL.

Zauschneria cana Greene
Epilobium canum (Greene) Raven ssp. canum

Yates ( 1 889) CR; Jepson ( 1 909-1 943) CR, AN; Abrams
and Ferris (1923-1960) CR, CA; Davidson and
Moxley ( 1 923) CR, AN; Munz ( 1 935) CR, CA; East-
wood(1941)CR, AN, CA; Dunkle (1942) AN; Dun-
kle (1950) RO, CR, CA; Munz and Keck ( 1 959) CR,
AN, CA; Thome (1967) CA; Munz (1974) CR, AN,
CA; Smith (1976) CR, AN.

Zauschneria villosa Greene
Epilobium canum (Greene) Raven ssp. canum

Greene (1887a) CR; Yates (1889) CR; Davidson and
Moxley (1923) RO, CL; Eastwood (1941) RO, CR,
CA, CL.

Zostera marina L. var. latifolia Morong
Zostera marina L.

Smith (1976) Channel Islands.

ACKNOWLEDGMENTS

I thank the curators of the following institutions for the loan
and on-site study of material for this paper: CAS-DS, CM,
F, GH, MO, ND-G, NY, RSA-POM, SBM, UC, UCSB,
UCSB-SCIR, US and the Pacific Missile Test Center at Point
Mugu, California. I greatly appreciate the written and oral
comments on certain critical groups, or areas, of the following
persons: D. Anderson ( Phalaris ), D. Barbe (weeds), R.M.
Beauchamp (San Clemente Island), S. Broich ( Lathyrus ), R.K.
Rmmmin (Calystegia), W.D. Clark ( Haplopappus ), W.S. Da-
vis ( Malacothrix ), A. Day ( Gilia ), L.T. Dempster (Galium),
D. Dunn ( Lupinus ), W. Ferren ( Suaeda ), L.D. Gottlieb
( Stephanomeria ), L. Heckard (Castilleja and Orobanche), P.
Hoch (Epilobium), J.S. Lassetter ( Vicia ), H. Lint (Juncus),
D.W. McNeal (Allium), R. Moran ( Dudleya , and Guadalupe
Island), N. Morin (Githopsis), R. Price (Erysimum), P. Raven
(Epilobium), A. Smith (ferns), C. Smith (northern islands),
D. Smith (Pityrogramma), L. Urbatsch (Haplopappus), D.H.
Wagner (Polystichum). I am very grateful to Dr. Robert
Thome for verifying several determinations and to S. Junak
for his determination of a report of Lomatium to be a lo-
cational error. I finally thank all those who encouraged this
project during its early stages, particularly Dr. Robert Thome.

Support from the California Arboretum Foundation, Nat-
ural History Museum of Los Angeles County Foundation,
and the Wrigley Memorial and Botanical Gardens is appre-
ciated. Fieldwork on the islands was only possible due to the
kind permission of the following: University of California at

Santa Barbara (Santa Cruz Island), Mr. A1 Vail and the Santa
Barbara Museum of Natural History (Santa Rosa Island),
U.S. Navy (San Nicolas and San Clemente Islands), and San-
ta Catalina Island Conservancy (Santa Catalina).

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Accepted 27 September 1984.

136 Contributions in Science, Number 365

Wallace: Vascular Plants of the Channel Islands

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RUGOSE CORALS (COELENTERATA, ANTHOZOA) FROM
THE LOWER PERMIAN MCCLOUD LIMESTONE AT
TOMBSTONE MOUNTAIN, NORTHERN CALIFORNIA

Edward C. Wilson1

ABSTRACT. A rugose coral faunule consisting of one solitary and
three massive species occurs in an Upper Wolfcampian Series stra-
tum of the McCloud Limestone at Tombstone Mountain, Shasta
County, California. It is the youngest coral fauna in the formation
and has species in common with older parts of the formation and
with Lower Permian formations of Oregon and Spitsbergen. Wex-
olina tombstonensis n. gen., n. sp. is a very large solitary coral.
Kleopatrina ( K .) svalbardense Fedorowski was described from the
Lower Permian (Sakmarian) Treskelodden Formation of Spitsber-
gen. Langenheimia klamathensis Wilson ranges lower in the for-
mation elsewhere. Petalaxis occidentalis (Merriam) was described
from the Lower Permian Coyote Butte Formation of Oregon. Coarse-
ly clastic sediments, algae, and the corals indicate a high-energy,
shallow-water, warm, marine paleoenvironment.

INTRODUCTION

Skinner and Wilde (1965) zoned the Lower Permian Mc-
Cloud Limestone of northern California into informal fu-
sulinid Zones A to H. Zones A to G are Wolfcampian and
Zone H is Leonardian (Skinner and Wilde, 1965; Wilde,
1971). Wilson (1982) described the corals of the formation
from Zones A to F. This paper reports the corals from
Zone G.

STRATIGRAPHY

A section was measured and intensively collected in the
McCloud Limestone at Tombstone Mountain, Shasta Coun-
ty, California (Fig. 1). It is 1 765 ft. (538 m) thick and consists
of a lower Zone A part overlain unconformably by Zone G
and Zone H rocks (Fig. 2). The Zone A rocks have a small
faunule of poorly preserved solitary and fasciculate rugose
corals and tabulate corals (loc. 6178). The Zone G rocks
contain the coral faunule reported in this paper (Iocs. 6181—
6183). The Zone H rocks have only rare, poorly preserved,
fasciculate (?) corals ( Heritsch loidesl).

The corals occur in beds cropping out between 865 and
880 ft. (264 and 267 m) above the base of the section. The
locality is within Zone G of Skinner and Wilde (1965) which

Contributions in Science, Number 366, pp. 1-11
Natural History Museum of Los Angeles County, 1985

they considered to be Late Wolfcampian and correlative with
the Coyote Butte Formation of central Oregon on the basis
of six shared fusulinid species.

The corals, except for Wexolina tombstonensis n. gen., n.
sp., which presently is unknown outside the type locality,
have noteworthy stratigraphic ranges and geographic distri-
butions. Kleopatrina (K.) svalbardense Fedorowski, 1980, was
described from the Lower Permian (Sakmarian) Treskelod-
den Formation of southern Spitsbergen. Its presence in north-
ern California suggests a correlation between the two areas
despite the intervening distance. The single specimen of Lan-
genheimia klamathensis Wilson, 1982, was originally de-
scribed from Zone F in the formation farther south. Its oc-
currence in Zone G at Tombstone Mountain establishes the
first stratigraphic range for the species (Fig. 3). Petalaxis
occidentalis (Merriam, 1942), erected for specimens from the
Lower Permian Coyote Butte Formation of central Oregon,
establishes a correlation of that formation with the McCloud
Limestone using corals. This corroborates Skinner and Wilde’s
(1965) fusulinid correlation.

Figure 3 incorporates ranges of the Zone G coral genera
into those from Zones A to F of the McCloud Limestone
previously reported by Wilson (1982).

Assignment of the McCloud Limestone coral faunas to the
Durhaminid Coral Province by Wilson (1982) is reinforced
by the discovery of Kleopatrina (K.) svalbardense, a species
previously known only from the Lower Permian of Spits-
bergen, also part of the Durhaminid Coral Province. Wex-
olina and Langenheimia, although presently unknown out-
side the McCloud Limestone, must be considered members
of the same province because of this association. Petalaxis,
widespread in the Durhaminid Coral Province, also occurs
in the westernmost Tethys Coral Province of Japan and pos-
sibly China (Sando, 1983).

1. Invertebrate Paleontology Section, Natural History Museum
of Los Angeles County, 900 Exposition Blvd., Los Angeles, Cali-
fornia 90007.

ISSN 0459-8113

Figure 1. McCloud Limestone exposures in Shasta County, California, showing locations of new measured section (7) at Tombstone Mountain
and other sections (1-6) from which corals previously were described by Wilson (1982).

2 Contributions in Science, Number 366

Wilson: Permian Corals of California

PALEOECOLOGY

The beds containing the corals described here, as well as the
whole of fusulinid Zone G at Tombstone Mountain, is a
coarse conglomerate with limestone clasts derived from all
the older McCloud Limestone fusulinid zones (Skinner and
Wilde, 1965). The corals described here occur in the matrix,
where the four species were found together. Reworked corals
in some of the clasts may be distinguished from the corals
in the matrix by careful examination although similar rock
colors and styolitic resorption makes this difficult in places.
The conglomerate indicates uplift of the older parts of the
McCloud Limestone and redeposition in a shallow, wave-
and current-affected, high-energy environment.

The presence of only cerioid colonial corals may represent
a response to turbulent water, in which massive shape is more
competent than a fasciculate shape. Similarly, the gigantic
size of the single solitary coral species may represent adap-
tation to strong wave action. The cerioid corals did not grow
to a very large size, perhaps because of rapid deposition. I
saw none with a diameter greater than 0.5 m and most were
a fraction of that. Coralla of the four species apparently lived
as a community among the cobbles and in the intervening
sandy stretches but generally not touching. Some were over-
turned prior to burial.

Random collecting of the cerioid coralla produced 40 spec-
imens, 27 of which are Petalaxis occidentalis, seven Kleo-
patrina (K.) svalbardense, and six Langenheimia klamath-
ensis. The first species clearly dominates numerically and its
presence elsewhere may indicate a comparable shallow-water,
high-energy environment. Intensive collecting yielded only
seven coralla of Wexolina tombstonensis, although I saw a
few others that could not be collected.

A few algal stromatolites indicate a paleoenvironment in
the euphotic zone. Sando (1980) considered algal-coral as-
sociations in Mississippian rocks to indicate a depositional
depth of less than 50 m. Wells (1957) considered colonial
rugose corals indicative of well-oxygenated, gently circulat-
ing, marine water with annual temperature minima of 16°
to 21°C. Most paleogeographic maps place the Permian equa-
tor on the west coast of North America somewhere in Cal-
ifornia or Baja California, suggesting a tropical environment
for the present site of the McCloud Limestone.

An earlier statement that “the McCloud Limestone was
deposited in clear, shallow, warm marine water with full
access to the open seas” (Wilson, 1982) is supported by the
paleoenvironmental evidence discussed above.

SYSTEMATIC PALEONTOLOGY

Morphological terminology is from Easton (1944) and Hill
(1981). Because I consider present family-level systematics
of most Permian rugose corals to be highly provisional, I
have not used them. The genera are separated into solitary
and cerioid groups, and the cerioid corals are arranged al-
phabetically.

Locality and type numbers refer to the Natural History
Museum of Los Angeles County, Invertebrate Paleontology

Section (acronym LACMIP) locality register and type cata-
logues. All specimens are deposited in LACMIP.

SOLITARY CORALS

Genus Wexolina new genus

DIAGNOSIS. Corallum solitary, large; calyx deep, steep-
walled, inverted cone-shaped; septa numerous, reaching from
corallite wall far into tabularium, thin in dissepimentarium,
dilate in tabularium in neanic stages, becoming thin in count-
er quadrants in ephebic stages; cardinal septum short; car-
dinal fossula closed, high on wall of calyx; dissepimentarium
wide, normal; tabellae of two zones, axial and periaxial; axial
tabellae concave upwards to cytose; periaxial tabellae gen-
erally horizontal.

TYPE SPECIES. Wexolina tombstonensis n. sp.

DISCUSSION. In transverse section, Wexolina some-
what resembles Bothrophyllum Trautschold, 1879, Pseudo-
timania Dobrolyubova and Kabakovich, 1948, and Timania
Stuckenberg, 1895. It may be distinguished from Bothro-
phyllum by its closed cardinal fossula and counter septum
of normal length. It may be distinguished from Pseudoti-
mania by its counter septum of normal length. It may be
distinguished from Timania by its counter septum of normal
length and absence of alar fossulae. In addition, none of these
genera has the deep, inverted cone-shaped calyx formed by
two zones of tabellae that characterizes Wexolina.

Until neanic stages are known for Wexolina, familial re-
lationships cannot be established.

ETYMOLOGY. The genus is named for Mr. John Bonnett
Wexo.

Wexolina tombstonensis new species

Figures 4, 5a-f

DIAGNOSIS. Because Wexolina is monotypic, the di-
agnoses of the genus and of the type species are identical.

EXTERNAL DESCRIPTION. Corallites solitary, cylin-
drical in ephebic stages, with apical angles of 40° in late neanic
stages (2.5 to 4 cm diameters), apparently slightly curved
with no abrupt angles, very large diameters to 10.3 cm, lengths
unknown but apparently large; epitheca with annual (?) con-
strictions 1.5 cm apart, a few minor constrictions between,
and fine growth lines; calices very deep (6 cm), sides sloping
very steeply (50° to 70°) downwards and inwards, lacking
central flat floor, cardinal fossulae prominent, located more
than half-way up calyx sides; axial boss absent.

TRANVERSE SECTION DESCRIPTION. Corallites
circular, 10.3 cm maximum diameter; septa of 2 orders, 84
to 88 each at maturity, all extending to corallite wall; major
septa thin in dissepimentarium, highly dilate in cardinal
quadrants in tabularium, thin to dilate in counter quadrants
in tabularium, adaxially attenuate, some sinuous, some
lengthened far into tabularium (amplexoid?), 30 to 40 mm
long in adult stages (corallite diameters 10 cm), with short-
ened cardinal septum (24 mm long in holotype); minor septa
thin, very short, confined to outer 2 to 3 rows of dissepiments
in adult stages, not seen in earlier stages (abraded corallites);

Contributions in Science, Number 366

Wilson: Permian Corals of California 3

EXPLANATION

CONGLOMERATE

LIMESTONE

INTRUSIVE

COVERED

Figure 2. Columnar section of the McCloud Limestone at Tomb-
stone Mountain, Shasta County, California. Locality numbers at
right of column show position of the only two coral faunas in the
section. Fauna of the higher locality is the subject of this paper.
Lower locality is mentioned in “Stratigraphy” section.

septal microstructure of fibers at right angle to sinuous dark,
center line; cardinal fossula closed, with neighboring several
pairs of septa arched around adaxial end, 32 mm deep in
holotype; tabularium filled with chaotic pattern of sections
of tabellae and attenuate septal ends; dissepimentarium wide,
width 15 to 20 mm in adult stages, not observed in younger
stages (abraded), lacking inner wall at tabularium interface;

dissepiments in 12 to 20 ranks in adult stage, regular pe-
ripherally, herringbone, angular, or straight adaxially; cor-
allite wall very thin, width 0.1 to 0.2 mm.

LONGITUDINAL SECTION DESCRIPTION. Disse-
pimentarium in adult stage (diameter 95 mm) of 19 to 22
ranks of steeply dipping dissepiments of very mixed sized:
generally small and globose near wall, becoming larger and
more elongate adaxially; tabellae of two zones, axial and
periaxial; axial tabellae of 10 to 12 ranks each side of ap-
proximate center of corallite (not indicated structurally), very
variable, most near center concave upwards, most near sides
steeply dipping downwards and inwards; periaxial tabellae
in 1 to 2 ranks, generally flat and horizontal, some cystose.

DOCUMENTATION. LACMIP holotype 7170, LAC-
MIP paratypes 7171-7176. Ten thin sections and 4 1 polished
sections from eight coralla from LACMIP localities 6181
(paratypes 7 1 72-7 1 74), 6 1 82 (holotype 7 1 70, paratype 7171),
and 6183 (paratypes 7175-7176) were studied. These three
localities are on strike within the same bed.

DISCUSSION. Most of the type specimens of this solitary
coral are preserved with the dissepimentaria mostly or wholly
removed by pre-burial abrasion. The calicular area of the
holotype is an exception but even it is progressively abraded
apically. The holotype has a dissepimentarium one-third the
diameter of the tabularium. Following is a list of sections
made of coralla from the type series, giving type designations,
corallite diameters, and septal numbers: holotype 7170, 10.3
cm — 88 septa, 9.7 cm — 84 septa; paratype 7171, 6.0 cm —
68 septa, 4.0 cm— 60 septa, 2.2 cm — 51 septa, 1.6 cm— 32
septa; paratype 7172, 3.5 cm — 55 septa; paratype 7173, 5.2
cm — 64 septa; paratype 7174, 5.0 cm — 67 septa, 3.5 cm —
52 septa; paratype 7175, 2.7 cm — 49 septa, 2.4 cm — 39 septa,
2.0 cm — 37 septa; paratype 7 1 76, 6.4 cm — 80 septa, 5.7 cm—
87 septa.

Unfortunately, the early ontogenic stages are unrepresent-
ed below 1.5 cm diameter eroded apices (probably originally
approximately 2.5 cm with dissepimentaria), although a spe-
cial search was made for small specimens. For this reason,
the genus and species cannot be assigned with certainty to
family.

The cyathopsid (?) of Wilson (1982:19, fig. 9f) was col-
lected from the McCloud Limestone at Potter Ridge, about
30 km south of Tombstone Mountain, where it occurs 38 m
below Langenheimia klamathensis. Specimens of the cy-
athopsid (?) are very poorly preserved, but figure 9f of Wilson
(1982) shows a corallite of similar large size to Wexolina
tombstonensis, somewhat more abundant septa (90), similar
sinuous septa protruding into the calyx, and a very similar
closed cardinal fossula with a short cardinal septum. Without
better preserved specimens it is impossible to tell if the cy-
athopsid (?) is referable to Wexolina, but these characters
suggest that it may be.

W. tombstonensis occurs in a conglomeratic unit at Tomb-
stone Mountain and is associated with abundant specimens
of three species (in three genera) of cerioid rugose corals.
These factors indicate that the environment of deposition,
and perhaps the living environment, was a very high energy
one. The preservation in the holotype of the calyx, the most

4 Contributions in Science, Number 366

Wilson: Permian Corals of California

WOLFC AMPI AN
FUSULINID
ZONES

ENIGMALITES

CLISIOPHYLLUM

c

D

HERITSC

HIOIDES

NEOMULTITHECOPORA

AULOPHY LLUM l?|

HETEROCANINIA

YATSENGI A

PETALAXIS

SYRINGOPORA

KLEOPATRINA I PORFIRIEVELLA I

MICHEUNtA

ARACHNASTRAEA

BAYH

AIUM

MCC

LOU Bl US

r

BASSIUS
KLEOPATRINA I KLEQPATRSNAj

DILLERIUM

TRASKINA

LANGENHE1MIA

DURHAMINA

©

WEXOLINA

©

Stratigraphic position uncertain but within Zone E or F

Figure 3. Stratigraphic ranges of corai genera in the McCloud Limestone in relation to Wolfcampian Series fusulinid zones of Skinner and
Wilde (1965). Zone G coral genera extensions reported in this study are added to previous ranges reported by Wilson (1982).

delicate part of the corallum, and the absence of the apical
parts beyond 10 cm below, where the corallum is badly erod-
ed, suggest that this corallum may have continued to live
and grow even after its apical parts were destroyed. Perhaps
coralla for this species lay loose on the substrate of such a
high energy environment that only the youngest parts nearest
the polyp remained intact. The highly dilate septa of the
tabularium may represent an adaption of a dense central core
resistant to erosion.

ETYMOLOGY. The species is named for Tombstone
Mountain.

CERIOID CORALS

Kleopatrina ( Kleopatrina ) svalbardense
Fedorowski, 1980

Figures 6a-d

Kleopatrina ( Kleopatrina ) svalbardense Fedorowski, 1980:
21, pi. 4, hgs. la-ld, 2, 3a-3b.

EXTERNAL DESCRIPTION. Corallum cerioid, hemi-
spheroidal, maximum observed diameter 14 cm; calyx not
observed; corallites parallel.

TRANSVERSE SECTION DESCRIPTION. Corallites 4
to 7 sided, 8 to 1 1 mm wide at greatest diameter; septa of 2
orders, 12 to 22 each, straight to slightly sinuous, rarely lons-
daleoid; major septa generally touching axial structure, es-
pecially cardinal and counter septa, 2.3 to 4 mm long, some-
what dilate in tabularium or thin throughout; minor septa
well developed, 1 to 2 mm long, generally crossing dissepi-
mentarium, extending into tabularium as nubs in some; dis-
sepimentarium generally regular, width 1.6 to 2.7 mm; dis-
sepiments concentric, herringbone, pseudoherringbone, or
lonsdaleoid (uncommon); axial structure clisiophylloid, cir-
cular to subcircular, rather small, 1 to 2 mm diameter, formed
of straight to slightly sinuous, somewhat thickened medial
plate connected to cardinal and counter septa, with radiating
straight to slightly sinuous septal lamellae touching axial ends
of many septa, connected by 2 to 4 axial tabellae; corallite
wall 0.1 to 0.2 mm wide.

Contributions in Science, Number 366

Wilson: Permian Corals of California 5

6 Contributions in Science, Number 366

Wilson: Permian Corals of California

Figure 4. Wexolina tombstonensis n. gen., n. sp. LACMIP holotype 7 1 70. Stereophoto of calyx. Cardinal fossula is near lower edge. Gastropod on left is Naticopsis

Figure 5. Wexolina tombstonensis n. gen., n. sp. a, b, LACMIP holotype 7170, transverse and longitudinal sections, c-f, transverse sections,
paratypes 7176 (c), 7171 (d-f). Partial transverse sections of Petalaxis occidentalis left of 5c and Langenheimia klamathensis left of 5d. x 1.

LONGITUDINAL SECTION DESCRIPTION. Dissep-
imentarium of 3 to 7 steeply to gently dipping ranks of me-
dium sized cystose (a few are elongate) dissepiments; tabellae
of 2 zones, axial and periaxial; periaxial tabellae generally
gently dipping in and up to axial tabellae, rarely flat, some
cystose, 1 5 to 22 per cm; axial tabellae in 1 to 3 ranks, large
to small, elongate to cystose, steeply sloping inwards and up
to medial plate, 15 to 23 per cm; some corallites lack axial
tabellae in places and have gently sloping tabulae instead.

DOCUMENTATION. LACMIP hypotype 7177-7178.
Three thin sections and 30 polished sections from hypotype
7177 from LACMIP locality 6181, three thin sections and
1 7 polished sections from one additional corallum from the
same locality, five thin sections and 53 polished sections from
two coralla from LACMIP locality 6182, two thin sections

and 23 polished sections from hypotype 7178 from LACMIP
locality 6183, and 3 1 polished sections from two coralla from
the same locality were studied.

DISCUSSION. This coral previously was known only from
the Lower Permian (Sakmarian) Treskelodden Formation of
southern Spitsbergen. Its occurrence in the Upper Wolfcamp-
ian of the McCloud Limestone provides an apparent bio-
stratigraphic correlation between the two formations despite
their great geographic separation.

Seven McCloud Limestone specimens are the basis for the
above description. Their characters fall within the range of
those of the type specimens from Spitsbergen, although some
coralla from the former occurrence have corallites a little
smaller in average diameters. The chief character distinguish-
ing this species from others in the genus is the nearly universal

Contributions in Science, Number 366

Wilson: Permian Corals of California 7

8 Contributions in Science, Number 366

Wilson: Permian Corals of California

Figure 7. Langenheimia klamathensis Wilson, 1982. LACMIP hypotype 7179, transverse (a) and longitudinal (b) sections. x3.

connection of the cardinal and counter septa to the complex
axial structure, which also is touched by other major septa,
giving it a distinct arachnoid appearance. The very short
septal-like structures on the external wall between major and
minor septa mentioned by Fedorowski (1980:21) can be seen
in some corallites of the McCloud Limestone specimens.

Langenheimia klamathensis Wilson, 1982

Figures 7a-b

Langenheimia klamathensis Wilson, 1982:65, figs. 37f,
38a-b.

Figure 6. Kleopatrina svalbardense Fedorowski, 1980. a, b, LACMIP hypotype 7177, transverse (a) and longitudinal (b) sections; c, d,
LACMIP hypotype 7178, transverse (c) and longitudinal (d) sections. x3.

Contributions in Science, Number 366

Wilson: Permian Corals of California 9

Figure 8. Petalaxis occidentalis (Merriam, 1942). LACMIP hypotype 7180, transverse (a) and longitudinal (b) sections. x3.

DESCRIPTION. The Tombstone Mountain specimens
referred to this species are so similar morphologically to the
type specimens from lower in the formation described by
Wilson (1982) that the original description will accommodate
them and is not repeated here.

DOCUMENTATION. LACMIP hypotype 7 1 79. Four thin
sections and 16 polished sections from the hypotype from
LACMIP locality 6181, two thin sections and 1 0 thin sections
from one additional corallum from the same locality, and
three thin sections and 20 polished sections from four coralla
from LACMIP locality 6182 were studied.

DISCUSSION. Occurrence of this species in fusulinid Zone
G at Tombstone Mountain extends its range upward in the
formation from the type locality in fusulinid Zone F at Potter
Ridge, about 30 km south of Tombstone Mountain. This
highly distinctive species is known only from these two lo-
calities.

Petalaxis occidentalis (Merriam, 1 942)

Figures 8a-b

Lithostrotion ( Lithostrolionella ) occidentalis Merriam, 1942:
377, pi. 56, figs. 2, 4, 7, 8, 11.

Petalaxis occidentalis (Merriam). Sando, 1983:32, pi. 20, figs.

1, 2.

DESCRIPTION. The Tombstone Mountain specimens
referred to this species are so similar morphologically to the
type specimens described by Merriam ( 1 942) that the original
description will accommodate them and need not be repeated
here.

DOCUMENTATION. LACMIP hypotype 7 1 80. Two thin
sections and 24 polished sections from the hypotype from
LACMIP locality 6181, four thin sections and 21 polished
sections from two additional coralla from the same locality.

10 Contributions in Science, Number 366

Wilson: Permian Corals of California

two thin sections and 26 polished sections from 7 coralla
from LACMIP locality 6182, and four thin sections and 60
polished sections from 17 coralla from LACMIP 6183 were
studied.

DISCUSSION. This is the first report of Petalaxis occi-
dentalis outside the type locality in the Coyote Butte For-
mation of central Oregon. Its presence in fusulinid Zone G
at Tombstone Mountain reinforces Skinner and Wilde’s (1965:
13) correlation of their Zone G of the McCloud Limestone
with the Coyote Butte Formation on the basis of six species
of fusulinids common to both.

LOCALITIES

The following localities are entered in the LACMIP locality
register. They are located in the McCloud Limestone, fusu-
linid Zone G, Upper Wolfcampian, on Tombstone Moun-
tain, Shasta County, California, in the northwest lA of the
southwest */* of the southeast 1 U of section 19, township 37
north, range 3 west, Mt. Diablo Meridian and Base Line, as
shown on the U.S. Geological Survey 1 5 minute topographic
quadrangle of Dunsmuir, California (1954, scale 1:62,500).

6181. South along strike from the base of LACMIP locality
6183 and east of the southern edge of the southeastern
peak. Conglomeratic limestone with large solitary cor-
als and abundant cerioid corals.

6182. Ten feet (3.05 m) stratigraphically higher, down the
dip slope from LACMIP locality 6181. Conglomeratic
gray limestone with enormous solitary corals and ce-
rioid corals.

6183. This locality is 865 to 880 ft. (263.6 to 268 m) above
the base of a measured section that begins on the west
face of the mountain at the exposed base of the
McCloud Limestone west of the northern peak, offsets
from that peak to the southwest peak, and extends
eastward over the southeast peak and along an east-
trending ridge to the base of the overlying Nosoni
Formation. Limestone, medium gray, weathers same,
conglomeratic, thin-bedded, with abundant cerioid
corals (some overturned), large solitary corals, fusu-
linids.

ACKNOWLEDGMENTS

I am grateful to Peter T. Gavin, Phillip G. Owen, and Eric
Scott for field assistance in 1982, 1983, and 1984 on the
arduous trips to Tombstone Mountain. Mr. John B. Wexo
generously financed the 1984 field trip. The Natural History
Museum of Los Angeles County and its Foundation sup-
ported this study in many ways. Dr. Winfield Henn of the
Shasta-Trinity National Forest provided collecting permits.
Photographs were prepared by Mr. Richard Meier and figures
are by Miss Mary Butler. Preparation of specimens was done
by me using facilities at LACMIP.

REFERENCES

Dobrolyubova, T. A., and N.V. Kabakovich. 1948. Neko-
torye predstaviteli Rugosa srednego i verkhnego Kar-
bona Podmoskovnogo basseyna. Akademiya Nauk
SSSR, Paleontologicheskii Institut, Trudy 14(2): 1-37,
pis. 1-16. (In Russian.)

Easton, W.H. 1944. Corals from the Chouteau and related
formations of the Mississippi valley region. Illinois State
Geological Survey, Report of Investigations 97:1-93.

Fedorowski, J. 1980. B. Corals. In Corals of the Treske-
lodden formation (Lower Permian) at Triasnuten, Horn-
sund, south Spitsbergen, ed. K. Birkenmajer, and J. Fe-
dorowski. Studia Geologica Polonica 66( 1 1 ):7— 27, pis.
1-6.

Hill, D. 1981. Rugosa and Tabulata. Part F. Coelenterata,
Supplement 1, vols. 1-2, 762 pp., 462 figs. In Treatise
on invertebrate paleontology, ed. R.C. Moore. Univer-
sity of Kansas Press, Lawrence.

Merriam, C.W. 1942. Carboniferous and Permian corals
from central Oregon. Journal of Paleontology 1 6(3):372—
81.

Sando, W.J. 1980. The paleoecology of Mississippian corals
in the western conterminous United States. Acta Pa-
laeontologica Polonica 25(3— 4):6 1 9—3 1 .

. 1983. Revision of Lithostrotionella (Coelenterata,

Rugosa) from the Carboniferous and Permian. U.S.
Geological Survey Professional Paper 1247:1-52, pis. 1-
20.

Skinner, J.W., and G.L. Wilde. 1965. Permian biostratig-
raphy and fusulinid faunas of the Shasta Lake area,
northern California. University of Kansas Paleontolog-
ical Contributions, Protozoa 6:1-98, pis. 1-65.

Stuckenberg, A. 1895. Korallen und Bryozoen der Stein-
kohlenablagerungen des Ural und des Timan. Geologi-
cheskii Komitet. Trudy ( Comite Geologique Memoires)
10(3): 1-244, pis. 1-24.

Trautschold, H. 1879. Die Kalkbruche von Mjatschkowa:
eine Monographic des obem Bergkalks. Societe Impe-
rial Naturalistes Moscou, Memoire 14:1-82, pis. 1-7.

Wells, J.W. 1957. Corals. Geological Society of America
Memoir 67(2):773-82.

Wilde, G.L. 1971. Phylogeny of Pseudofusulinella and its
bearing on Early Permian stratigraphy. Smithsonian
Contributions to Paleobiology 3:363-79.

Wilson, E.C. 1982. Wolfcampian rugose and tabulate cor-
als (Coelenterata: Anthozoa) from the Lower Permian
McCloud Limestone of northern California. Contribu-
tions in Science, no. 337, 90 pp. Natural History Mu-
seum of Los Angeles County.

Accepted 1 April 1985.

Contributions in Science, Number 366

Wilson: Permian Corals of California 1 1

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Number 367
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CONTRIBUTI

THE LATE MIOCENE DOLPHIN PIT HA NODELPHIS ABEL,

1905 (CETACEA: KENTRIODONTIDAE)

FROM CALIFORNIA

Lawrence G. Barnes

B

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Edward C. Wilson

Printed at Ailen Press, Inc . Lawrence, Kansas

THE LATE MIOCENE DOLPHIN PIT HA NODELPHIS ABEL,
1905 (CETACEA: KENTRIODONTIDAE)

FROM CALIFORNIA

Lawrence G. Barnes1

ABSTRACT. Fossil species of the unusual and relatively highly
derived kentriodontid dolphin genus Pithanodelphis Abel, 1905, are
now known from rocks bordering both the North Atlantic and the
North Pacific Ocean basins. The type species of this genus, Late
Miocene Pithanodelphis cornuius (du Bus, 1872), is known from the
Antwerp Basin in Belgium. A skull illustrated by Abel in 1905 is
here designated as the iectotype of this species. An approximately
contemporaneous species, Pithanodelphis nasalis, new species, dif-
fers from P. cornutus by having a different suite of derived and
primitive cranial characters and it is one of the most abundant fossil
cetaceans in the Late Miocene age Monterey and Modelo formations
in southern California.

Bones around the nares and cranial vertex of species of Pithano-
delphis are asymmetrical, but in a unique manner that is totally
unlike the condition in species of modem Delphinidae and other
Odontoceti which possess cranial asymmetry'. The nasal bones are
very large. In recognition of these derived characters, Pithanodelphis
is classified in a new' subfamily of Kentriodontidae, the Pithano-
"> delphinae. Apparently Pithanodelphis nasalis was able to produce
sounds used for echolocatkm, lived in moderately deep water off-
shore over the continental shelf, and had a heterogeneous diet com-
prised mostly of small fishes. Adult individuals attained a body
length of approximately 200 cm. The vertebral column is similar to
those of the Middle Miocene fossil kentriodontine dolphin, Delphin-
odon dividum True, 1912b, and the extant bottlenosed dolphin, T ur-
siops truncaius Montagu, 1821.

INTRODUCTION

Recent studies have shown that during Miocene time an
extinct group of dolphins, the family Kentriodontidae, which
was first recognized from fossils found in deposits around
the North Atlantic Ocean, also had an equally significant
history in the North Pacific Ocean (Bames, 1977, 1978; Barnes
and Mitchell, 1984). Five kentriodontid genera are now known
to have had species that lived in both ocean basins, and one
of these is the genus Pithanodelphis Abel, 1905. The type

Contributions in Science, Number 367, pp. 1-27
Natural History Museum of Los Angeles County, 1985

species of the genus is P. cornutus (du Bus, 1872), which is
known only from specimens found in Late Miocene sedi-
ments in the Antwerp Basin in Belgium. I have previously
(Bames, 1977) recognized the genus Pithanodelphis from the
approximately correlative Monterey Formation in southern
California, and classified the genus in the nominate kentri-
odontid subfamily Kentriodontinae (Bames, 1978).

The fossils from California represent a new species that I
described in a Ph.D. dissertation (Bames, 1 972). This species
is known by several skulls and some postcranial bones, most
of which are from a vertically and laterally restricted strati-
graphic section of the Monterey Formation in the San Joa-
quin Hills in Orange County, California, near the southern
limit of the topographic and depositional feature known as
the Los Angeles Basin (Woodford et al., 1954). Some of the
fossils were discovered in naturally formed outcrops, but
most were discovered in 1969 as a result of earth-moving
for construction of an extensive commercial/manufacturing
complex for North American Rockwell Land Corporation
near Aliso Creek in the Laguna Niguel district, part of which
later became the offices of the United States General Services
Administration. This site has been discussed by Domning
(1978), Howard (1978), and Bames, Raschke, and McLeod
(in press). One additional specimen that I refer to this species
is from the correlative Upper Member of the Modelo For-
mation that is exposed 84 km to the north of Laguna Niguel
in the Santa Monica Mountains, Los Angeles County, Cal-
ifornia. The purpose of this paper is to diagnose these fossils
as a new species and to describe the available sample of
specimens.

1 . Section of Vertebrate Paleontology, Natural History Museum
of Los Angeles County, Los Angeles, California 90007.

ISSN 0459-8113

METHODS AND MATERIALS

In this study, the closest comparisons are made between the
new species and two other kentriodontids. I refer the reader
to descriptions and illustrations of specimens of Kentriodon
pernix Kellogg, 1927, provided by Kellogg (1927, 1928),
Barnes (1978, including other species of kentriodontids), and
Barnes and Mitchell (1984), and of Delphinodon dividum
T rue, 1912b, provided by T rue (1912b), Barwick (1939), and
Barnes (1978). I have also made comparisons with various
living odontocete taxa. Systematics and ranges of these are
provided by Hershkovitz (1966) and Rice (1984).

Measurements of skulls and the mandible were made fol-
lowing the methods outlined by Perrin (1975). In the tables
of measurements, a number in parentheses following the de-
scription of a parameter refers to the same measurement of
Perrin. Anatomical terminology is derived from Kellogg
(1927), Fraser and Purves (1960), Kasuya (1973), Barnes
(1978, 1984), and Barnes and Mitchell (1984). The repository
of specimens, the Natural History Museum of Los Angeles
County, Los Angeles, California, is abbreviated LACM. Pre-
cise locality descriptions may be provided to qualified in-
vestigators upon request.

In the illustrations, bones and other anatomical structures
are labeled according to the following abbreviations:

aon— antorbital notch

Bo— basioccipital bone

Bs— basisphenoid bone

ch— cranial hiatus

earn— external acoustic meatus

fio— ventral apertures, infraorbital foramen

fmx — maxillary foramen

fop— optic foramen

fpal— palatine foramen

fpmx— premaxillary foramen

Fr— frontal bone

gf— glenoid fossa

Ju— jugal bone

La — lacrimal bone

Met— mesethmoid bone

mrg— mesorostral gutter

ms— middle sinus

Mx— maxillary bone

n— naris

Na— nasal bone

Oc— occipital bone

occ— occipital condyle

Pa— parietal bone

Pal— palatine bone

Pm x— premaxillary bone

pop— paroccipital process

Pt— pterygoid bone

Pt(ll) — lateral lamina of pterygoid

Pt(ml)— medial lamina of pterygoid

pts— fossa for pterygoid sinus

sq — squamosal bone

sqf— squamosal fossa
Vo— vomer bone

SYSTEMATICS

Class Mammalia Linnaeus, 1758
Order Cetacea Brisson, 1762
Suborder Odontoceti Flower, 1867

Superfamily Delphinoidea (Gray, 1821)
Flower, 1864

Family Kentriodontidae (Slijper, 1936)
Barnes, 1978

Kentriodontinae Slijper, 1936:556; as a subfamily of the fam-
ily Delphinidae.

Kentriodontidae. Slijper, 1 958:label in fig. 36; emended rank
without explanation in text.

Kentriodontidae. Barnes, 1978:3; emended rank, as a family
of the superfamily Delphinoidea.

EMENDED DIAGNOSIS OF FAMILY. A family in the
superfamily Delphinoidea differing from Albireonidae, Mon-
odontidae, Delphinidae, and Phocoenidae by having skulls
with small pterygoid sinus in pterygoid hamulus; differing
from Monodontidae, Delphinidae, and Phocoenidae by lack-
ing excavation in exoccipital for posterior sinus and in lateral
side of basioccipital for peribullary and/or pterygoid sinus,
by having symmetrical posterior ends of premaxillae which
contact nasals on both right and left sides, and by having
symmetrical cranial vertex (except in case of kentriodontids
with asymmetrical vertex, which have midline between na-
sals twisted to right instead of to left); differing from Del-
phinidae and Phocoenidae by lacking anterior sinus and lack-
ing large posterodorsal extension of antorbital lobe of
pterygoid sinus between frontal and maxilla; differing from
Albireonidae and Phocoenidae by lacking premaxillary em-
inences and having instead, wide, flat, and elevated spiracular
plates on premaxillae on either side of external nares; dif-
fering from Monodontidae and Delphinidae by having sym-
metrical mesethmoid and external nares, equal areas of ver-
tex covered by right and left nasals, and by having right and
left spiracular plates approximately equal in size; and differ-
ing from Albireonidae by having mesorostral gutter open
dorsally.

INCLUDED SUBFAMILIES. Kampholophinae Barnes,
1978; Kentriodontinae (Slijper, 1936) Barnes, 1978; Lopho-
cetinae Barnes, 1978; and Pithanodelphinae, new subfamily.

Subfamily Pithanodelphinae, new subfamily

DIAGNOSIS OF SUBFAMILY. A subfamily of Ken-
triodontidae differing from Kampholophinae, Lophocetinae,
and Kentriodontinae by having skulls with asymmetrical cra-
nial vertex in which midline between nasals bends toward
right side posteriorly, right maxilla encroaches farther than
left toward midline posteriorly, right spiracular plate is slight-

2 Contributions in Science, Number 367

Barnes: Pithanodelphis from California

ly wider than left, and right nasal is higher than left, posterior
end of premaxilla extending as slender projection between
nasal and maxilla rather than wide, abruptly terminating and
with elevated posterolateral comer, nasal bone much larger
and very convex, olfactory fontanelle present in posterior
wall of each naris; differing further from Kampholophinae
and Kentriodontinae by having periotic with narrower, more
transversely compressed anterior process; and differing fur-
ther from Kentriodontinae by lacking obliquely oriented sul-
cus on anterolateral surface of nasal bone within upper part
of naris.

TYPE AND ONLY INCLUDED GENUS. Pithanodel-
phis Abel, 1905, Late Miocene, Belgium and California.

Pithanodelphis Abel, 1905

Phocaenopsis Huxley, 1859, part, du Bus, 1872:500.
Pithanodelphis Abel, 1905:142; for Phocaenopsis cornutus

du Bus, 1872, only.

EMENDED DIAGNOSIS OF GENUS. The same as for
the subfamily until other genera are assigned to the subfam-
ily.

TYPE SPECIES. Pithanodelphis cornutus (du Bus, 1872).

INCLUDED SPECIES. Pithanodelphis cornutus (du Bus,
1872); and Pithanodelphis nasalis, new species.

Pithanodelphis nasalis, new species

Figures 1-14

aff. Pithanodelphis Abel, 1905. Bames, 1977:328 (table 4).

DIAGNOSIS OF SPECIES. A species of Pithanodelphis
differing from P. cornutus by having skull with more prom-
inent lambdoidal and occipital crests, occipital shield smaller
and not as convex, temporal fossa curving farther around
margin of occipital shield, nasal relatively larger, zygomatic
process of squamosal smaller with more tapered anterior end.

HOLOTYPE. LACM 30093, a nearly complete skull, dor-
soventrally crushed, lacking much of the basicranium, bear-
ing 23 whole or incomplete teeth in place, with 16 loose
teeth, right and left tympanic bullae, left periotic, malleus,
incus, and stapes; right and left dentaries with 1 1 whole or
partial teeth in place, collected by W. Earl Calhoun and Mi-
chael K. Hammer in 1969.

TYPE LOCALITY. LACM locality 5077, Laguna Niguel
district, San Joaquin Hills, Orange County, California.

PARATYPES. LACM 26635, an undistorted but badly
shattered incomplete skull lacking anterior end of the rostrum
and basicranium, bearing five whole or partial teeth in place,
and one fragment of the extremity of the rostrum, collected
at LACM locality 5069 by W. Earl Calhoun in 1969. LACM
29087, two fragments of a skull (the dorsolateral comer of
the braincase and the left squamosal), partial vertebral col-
umn and ribs, collected at LACM locality 5082 by Marion
J. Bohreer and W. Earl Calhoun in 1969.

REFERRED SPECIMENS FROM THE MONTEREY
FORMATION. LACM 31186, right side of rostrum col-

lected at LACM locality 507 1 by Marion J. Bohreer in 1969;
LACM 1 22670, left periotic collected by Michael D. Quarles,
8 November 1982, and LACM 123872, skull, collected by
David P. Whistler and L.G. Barnes, 31 May 1975, both from
LACM locality 6902; LACM 123873, skull collected at LACM
locality 1101 by Michael K. Hammer.

FORMATION AND AGE. The holotype, paratypes, and
referred specimens listed above are all from rocks that have
been identified as the upper part of the Monterey Formation
(Vedder, Yerkes, and Schoellhamer, 1957; Fife, 1974), cor-
related with the “Margaritan” provisional mega-invertebrate
stage of Addicott (1972), with the Mohnian foraminiferal
stage, and indirectly with the later part of the Clarendonian
North American land mammal age, and are of Late Miocene
age, approximately 10 to 11 million years old (Bames, 1977;
Repenning and Tedford, 1977; Howard, 1978). This part of
the Monterey Formation has yielded a diverse fossil verte-
brate assemblage (Bames, Raschke, and McLeod, in press),
including the bird fossils that were described by Howard
(1976, 1978), the sirenian fossils reported by Domning(!978),
and a fairly diverse cetacean assemblage reported by Bames
(1977). Stratigraphically lower within the Monterey For-
mation in the same district of Orange County vertebrate
fossils, including birds (Howard, 1966, 1968) and cetaceans
(Bames, 1978), have been reported that have closer affinities
with Middle Miocene assemblages in California, especially
with the Sharktooth Hill Local Fauna in central California.
These older assemblages in the Monterey Formation have
been correlated indirectly with the earlier part of the Clar-
endonian land mammal age (Howard, 1978:24).

In the upper part of the Monterey Formation in Laguna
Niguel, Pithanodelphis nasalis is associated with the gannet,
Morns lompocanus (Miller, 1925); the booby, IMiosula me-
dia Miller, 1925; the sea cow, Dusisiren jordani (Kellogg,
1 925a); and the pinnipeds, Pithanotaria starri Kellogg, 1 925b,
and Imagotaria downsi Mitchell, 1968, all of which were
originally based on specimens from the Late Miocene age
diatomites of the Sisquoc Formation near Lompoc, Santa
Barbara County, California. This association further rein-
forces the age yielded by the correlations given above (see
also Repenning and Tedford, 1977).

Because the holotype and paratypes of Pithanodelphis na-
salis were exposed during a construction project as men-
tioned in the introduction, they were collected in a salvage
situation and the precise stratigraphic relationships between
them were not recorded. The holotype, LACM 30093, was
collected from a fine-grained yellow siltstone bed that was
approximately 6 to 8 inches thick and was exposed within a
section of otherwise fairly uniformly bedded white diatomite
1 40 m north of the excavation for the foundation of the main
North American Rockwell building. The paratype skull,
LACM 26635, was found in a loose, coarse-grained, bed of
gray sand uncovered near the northeast comer of the same
building. Because the strata in this immediate area generally
dip toward the north, the bed that yielded the paratype skull
was probably stratigraphically lower in the Monterey For-
mation than the one that yielded the holotype, but was prob-
ably no more than 10 m lower.

Contributions in Science, Number 367

Barnes: Pithanodelphis from California 3

The paratype partial skeleton, LACM 29087, was collected
from a coarse sand bed within bedded diatomite at the north-
west comer of the building, and its stratigraphic position was
therefore probably also lower than the holotype. The ver-
tebral column of the paratype was bisected in the thoracic
area by machinery that was cutting a trench, and Bohreer
and Calhoun independently collected opposite ends of the
skeleton. After later conversations with both men I was con-
vinced that only one individual fossil skeleton was involved.

Another partial skull, LACM 123872, and an isolated peri-
otic, LACM 122670, referred to the species were both col-
lected near the base of a very coarse, ca. 3-m-thick bed of
yellow sand exposed in a road cut 400 m northwest of the
main building site. The sand bed that yielded these fossils
has subsequently been observed in the field to lie stratigraph-
ically above the diatomite that produced the holotype, and
it is the highest part of the Monterey Formation that is ex-
posed in the vicinity (Barnes, Raschke, and McLeod, in press).

The referred skull, LACM 1 23873, is from another nearby
site, LACM 1101, the same locality that yielded the partial
skeleton of an otariid pinniped (LACM 1404) that was iden-
tified by Downs (1955) as cf. Allodesmus kernensis Kellogg,
1922. This specimen is actually an imagotariine otariid, pos-
sibly Imagotaria downsi or a closely related species. I. downsi
is known from such Late Miocene (Clarendonian age) for-
mations in California as the Towsley, Sisquoc, and Santa
Margarita (Repenning and Tedford, 1977). The locality
(LACM 1 101) is 1.4 km from the other Laguna Niguel sites
mentioned above that yielded Pithanodelphis nasalis, and is
separated from them by the wide valley that was formed by
Aliso Creek and, thus, no direct stratigraphic correlation is
possible. The site is, however, in a well-bedded diatomite
very much like that which yielded the holotype skull, and
this diatomite is overlain by a coarse yellow sand bed like
that which produced the referred specimens of P. nasalis at
LACM locality 6902. The strata are undoubtedly correlative,
and I therefore conclude that all the specimens from these
Laguna Niguel localities were collected within the uppermost
few tens of meters of the Monterey Formation.

REFERRED SPECIMEN FROM THE MODELO
FORMATION. LACM 15196, incomplete skull and jaws
with some associated postcranial bones collected from LACM
locality 1230, Studio City, Los Angeles County, California,
by Terry and Michael Pohl about 1955.

FORMATION AND AGE. Upper Member of the Modelo
Formation, Late Miocene age, correlated with the upper part
of the Monterey Formation in the San Joaquin Hills (Wood-
ford et al., 1954:fig. 2). The locality is within outcrops of
diatomaceous shale that have been mapped as the Upper
Member of the Modelo Formation of Late Miocene age
(Hoots, 1931). A nearby outcrop of the same rock unit pro-
duced the fossil bird, Sula pohli Howard, 1958.

ETYMOLOGY. The species name, nasalis, is derived from
Latin, nasus, for nose, and is in reference to the exceptionally
large nasal bones of this species.

DESCRIPTION. Skull. The description and reconstruc-
tions (Figs. 3, 5, 8) of the skull of Pithanodelphis nasalis are
composites, being based on all the available skulls. The ho-

lotype is the most complete known skull, but unfortunately
it is also the most distorted, the braincase being flattened
with the basicranium being pushed to the left side. The para-
type skull, LACM 26635, although badly shattered, was not
distorted by sediment compaction and exhibits the original
proportions of the braincase, thus providing information on
the undistorted facial surface and true cranium height. In-
formation on the structure of the zygomatic process, rostrum,
and palate was obtained mostly from the holotype (LACM
30093). Both the paratype and holotype yielded data on the
squamosal and pterygoid regions. Both of these skulls con-
firmed the confident identification of the paratype partial
skeleton, which includes two skull fragments. The referred
skull, LACM 1 5 1 96, from the Modelo Formation, is the only
one with the occipital condyles intact. My restoration of the
shape of the pterygoid hamulus and its contained fossa for
an air sinus is tenuous. The holotype is crushed obliquely in
this area, but it preserves the shapes of the pterygoid-palatine
suture, the posterior opening of the sinus and the semicircular
notch in the lateral lamina of the pterygoid, and shows that
the anterior part of the pterygoid sinus is virtually the same
width as the posterior part, and that the hamulus is contin-
uously floored by thin bone. The paratype and holotype skulls
both show the relationships between the pterygoid and pal-
atine, and the exposure of the vomer between the pterygoid
hamuli.

The sample of skulls includes a range of sizes, with the
holotype near the mean (Table 1). The paratype skull, LACM
26635, is the largest and has osteological characters indica-
tive of advanced maturity: its crests, tuberosities and nasal
bones are prominent.

Characters that indicate that the small skull referred to the
species, LACM 123872, is from a juvenile individual are:
small size, short postorbital process of the frontal, incom-
pletely formed vomer on the palate between the pterygoids
and a cleft between the frontals on the cranial vertex.

Pithanodelphis nasalis has skull proportions like the living
freshwater South American stenine delphinids in the genus
Sotalia Gray, 1866. The facial surface is roughly square, the
rostrum is narrow and of medium length with a broad base,
and the braincase is highly vaulted with prominent comers,
widely flaring zygomatic processes, and a high vertex.

The premaxillae occupy most of the dorsal rostral surface,
have dense surface structure, and on the holotype extend
only 7 mm anteriorly beyond the maxillae. They are not
fused medially but are closely appressed over the midline of
the mesorostral gutter at rostral midlength. The rostral part
of each premaxilla is transversely convex, becoming flat-lying
posteriorly and nearly vertically oriented at the anterior end.
The mid-part of each premaxilla is depressed around the
premaxillary foramen, but the medial edge next to the me-
sorostral gutter is elevated. The premaxillary foramina are
situated slightly anterior to the location of the antorbital
notches and approximately equidistant between the medial
and lateral premaxillary margins. A faint anteromedial sulcus
extends anteriorly from each premaxillary foramen and con-
verges toward the medial margin, intersecting it at a point
nearly one-third of the distance to the anterior end of the

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Barnes: Pithanodelphis from California

rostrum. Medial to this faint groove, the surface of the pre-
maxilla is rugose, and this marks the area of attachment of
the nasal plug muscle (Lawrence and Schevill, 1956).

A deeper posterolateral sulcus extends posteriorly from
each premaxillary foramen, toward the lateral margin of the
premaxilla, reaching it at a point over the middle of the orbit.
There is only a very faint posteromedial sulcus. Adjacent to
the lateral narial margins the spiracular plates are broad,
elevated, and convex. The posterior terminations of the pre-
maxillae narrow abruptly and have a slender posterior pro-
jection that is constricted between the maxillae and the swol-
len nasals. This thin projection is only 1 to 2 mm wide and
extends about half the remaining distance posteriorly to the
occipital crest between the maxilla and nasal.

On the anterior part of the rostrum the lateral surface of
the maxilla has a porous texture and presents a nearly vertical
surface adjacent to the alveolar row. The maxilla is thicker
posteriorly where it forms a squared margin of the rostrum
anterior to the antorbital notch and has only a narrow dorsal
surface exposure adjacent to the premaxilla. There are three
anterior maxillary foramina around the antorbital notches
and above the anterior part of the orbit, and one larger pos-
terior maxillary foramen over the posterior part of the orbit.
Two or three small foramina exit from the maxillary-pre-
maxillary suture anterior to the antorbital notch.

On the antorbital process the dorsal surface of the maxilla
is thin but rugose. Elsewhere on the facial surface the maxilla
is smooth, and its margins are elevated medially adjacent to
the nasal and posteriorly along the occipital crest. The pos-
teromedial comer of each maxilla extends around the pos-
terior side of the corresponding nasal bone to encroach on
the frontal where it is exposed at the cranial vertex. Qn all
of the skulls, the left maxilla does not extend medially as far
as does the right one. On the surface of each maxilla is a low,
crescent-shaped ridge extending from the area of the pos-
terolateral side of the nasal toward the temporal fossa. Such
ridges undoubtedly mark the attachment of one or more of
the layers of the nasal musculature (see Mead, 1975), and
are most pronounced on the most mature skull (LACM
26635).

The external nares are narrow anteriorly, wide posteriorly,
pass vertically into the skull and are separated by a relatively
high, thin mesethmoid septum. The right and left narial pas-
sages are equal in size and shape. A circular olfactory fora-
men, approximately 5 mm in diameter in LACM 26635, is
located in the center of the posterior wall of each naris. These
foramina are the vestiges of a more primitive condition in
which the olfactory lobe of the brain had a major connection
with the nasal passages. In some primitive fossil odontocetes
(e.g., ZarhachisC ope, 1868; Squalodon Grateloup, 1840;ylr-
gyrocetus Lydekker, 1894; Eurhinodelphis du Bus, 1867) the
mesethmoid and ectethmoid bones are distinct and divide
the primitively single olfactory fontanelle into two apertures.
In Recent delphinids the mesethmoid and ectethmoids are
fused into one solid plate and only tiny perforations remain
in some individuals and taxa to represent the olfactory fo-
ramina (see Kellogg, 1928:199-202). Pithanodelphis nasalis
demonstrates an intermediate condition in which the mes-

ethmoid and ectethmoids are fused but the olfactory foram-
ina are still relatively large.

There is no prominent suture between the mesethmoid
and nasal bones, and the posterior walls of the narial passages
merge smoothly with the vertical anterior surfaces of the very
bulbous nasals. A large, basin-like depression is on the mid-
line of the anterodorsal surfaces of the nasals. The nasals are
separated posteriorly by two vertical, median wedges of the
frontals. The frontals are also exposed on the vertex behind
the nasals, where they extend as wedges between the nasals
and the maxillae, and between the maxillae and the supra-
occipital. The nasal bones are proportionally larger than in
any other known delphinoid species. On all specimens the
left nasal is slightly lower and wider than the right. The suture
separating the two nasals bends to the right of the midline
of the braincase posteriorly, and the posterior ends of the
nasals, therefore, are shifted slightly to the right side of the
skull. In derived living odontocetes with asymmetrical cra-
nial vertices, the displacement of the nasal bones is always
to the left.

The occipital shield has an unusual conformation for a
delphinoid (Fig. 10). The occipital condyles, preserved only
on LACM 1 5196, are separated by a notch ventrally and are
relatively small for the skull size (Fig. 8). The occipital crest
and the laterally located lambdoidal crests are large and flare
posteriorly to outline the occipital shield, which is inclined
anterodorsally, has a generally convex surface, and bears a
sulcus dorsal to the foramen magnum. The temporal fossae
wrap posteriorly around the occipital shield to such an extent
that there is no more than a 50 to 60 mm distance between
the right and left lamboidal crests across the posterior surface
of the cranium (Fig. 10). The exoccipital is thin, vertically
oriented, and flares anterolaterally where it is appressed to
the posterior surface of the squamosal. On no specimen is
the paroccipital process complete. The posterior side of the
right maxilla protrudes farther posteriorly and therefore
pushes the occipital shield further posteriorly than does the
left maxilla.

The ventral surface of the rostrum is slightly up-curved
anteriorly and the palatal surface is generally flat except where
it becomes slightly convex posteriorly near the palatine bones.
The extent of palatal exposure of the vomer between the
maxillae varies in the sample. On the holotype it is about 35
mm long, but on the referred skull LACM 123872 it is 50
mm long. At the anterior end of the exposed vomer the
premaxillae appear on the palate medial to the maxillae. They
increase in width anteriorly, so that within a distance of 55
mm, the maxillae are entirely excluded from between the
alveolar rows.

The largest teeth are in the middle part of the tooth row,
where the alveoli are about 5 mm in diameter, and the an-
terior and posterior ones are progressively smaller. The al-
veoli are circular with the bone around them raised and
rugose. At the anterior rostral extremity, the alveolar rows
are very close together. They gradually diverge posteriorly
until they are 52 to 55 mm apart at their posterior ends. In
specimens with complete tooth rows, the number of upper
teeth on each side varies from 27 to 30. In the available

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Barnes: Pithanodelphis from California 5

6 Contributions in Science, Number 367

Barnes: Pithanodelphis from California

Figure 1. Pithanodelphis nasalis, new species, holotype, skull, LACM 30093. LACM locality 5077, dorsal view.

Contributions in Science, Number 367

Barnes: Pithanodelphis from California 7

Figure 2. Pithanodelphis nasalis, new species, paratype, skull LACM 26635, LACM locality 5069, dorsal view.

8 Contributions in Science, Number 367

Barnes: Pithanodelphis from California

fmx

Figure 3. Pithanodelphis nasalis, new species, composite reconstruction of skull, dorsal view, based on available specimens; for explanation of abbreviations see
Methods and Materials.

sample, the posterior part of the dentition is variable in the
following ways. The alveolar row stops at variable distances
from the antorbital notch, and this accounts for the above-
mentioned variation in the tooth count. In the holotype,
LACM 30093, the posteriormost alveoli are significantly
smaller than in the paratype, LACM 26635, and in the re-
ferred skull, LACM 123872. Some of the specimens show
crowding of the posterior teeth, so that in some cases two
adjacent alveoli merge.

A palatine foramen in each maxilla near the exposed vomer
is confluent with a groove that extends anteriorly and con-
verges on the midline. A larger, less well defined groove on
each maxilla originates at the maxillary-palatine suture and
extends anteriorly to converge on the midline of the palate.
The palatine bones, marked by arc-shaped sutures, extend
anteriorly on the palate to a point varying between 14 and
19 mm anterior to the level of the antorbital notches. The
maxillae extend posteriorly between the palatines at the mid-
line, between the pterygoid sinus fossae. On the paratype
skull (LACM 26635) the vomer descends between the pal-
atines to form a thin, deep keel that extends posteriorly be-
tween the nares. The palatines form the sides and roof of the
fossae for the pterygoid sinuses, which are shallow, triangular,
and pointed anteriorly. Parts of the thin, non-porous pter-
ygoid bone which floored the sinuses are preserved on the
holotype skull (LACM 29087). In this area, the bone is very
crushed but I can discern a hamular process and a semicir-
cular notch in the lateral lamina of the pterygoid.

The vomer forms a very thin keel separating the nares and
it underlies the basisphenoid bone as a thin horizontal plate,
extending posteriorly as far as the cranial hiatus. Posterior
to this point the basioccipital crest is broken away on all
specimens.

Within the orbit there are several foramina. A foramen,
elliptical in outline, pierces the lateral wall of the narial pas-
sage in the paratype (LACM 26635) and connects with the
orbit. The infraorbital foramen is in a position typical of
species of Recent Delphinidae— medial to the antorbital notch
and near the margin of the pterygoid; but it is smaller, sim-
pler, and not surrounded by struts of bone. The orbital ap-
erture of the foramen is 4 mm in diameter and connects with
a branch extending anteriorly within the rostrum to emerge
as the premaxillary foramen. Lateral to this, but still con-
fluent with the infraorbital foramen, a smaller foramen con-
nects to the anterior maxillary foramina, and posteriorly
another branch leads to the posterior maxillary foramen. A
shallow fossa is excavated in the ventral surface of the frontal
anterior to, and separated from, the tract of the optic nerve
by a thin, elevated crest. This fossa was undoubtedly the
location of a small preorbital lobe of the pterygoid air sinus.
The sinus did not extend dorsally between the frontal and
the maxilla as it does in phocoenids, and, to a lesser extent,
in some Recent species of Delphinidae. Another fossa in the
frontal bone, posterior to the tract for the optic nerve,
marks the site of a postorbital lobe of the pterygoid air sinus
(Fraser and Purves, 1960).

The orbit is relatively smaller than in a delphinid such as
the Recent common dolphin, Delphinus delphis Linnaeus,

1758, but it is still relatively large for a kentriodontid. The
antorbital process of the frontal is prominent. The lacrimal
bone comprises the anteroventral surface of the antorbital
process and its anterolateral part is very thick, but it becomes
thin and narrow medially. The lacrimal is wedged into a
shallow depression in the ventral surface of the maxilla me-
dial to the antorbital notch. The jugal is fused to the anterior
edge of the lacrimal and protrudes into the middle of the
antorbital notch, thereby forming a small eminence. This
eminence is particularly well preserved on the right side of
the holotype (LACM 30093). Such an eminence is unusual
for a delphinoid, but a similar one occurs in some species of
Recent beaked whales (family Ziphiidae).

The left jugal of the holotype is entirely preserved and
appears to retain its approximate original curvature. It mea-
sures approximately 80 mm in curvilinear length, is slender
and round in cross section anteriorly, and is flattened trans-
versely in its posterior part where it contacts the zygomatic
process of the squamosal. It articulates with the squamosal
on a small, anteriorly directed process on the ventral margin
of the zygomatic process that is in a position similar to that
in Delphinus delphis.

The temporal fossa is very large, elongate and expanded
dorsoventrally. Its anterior part is overhung by the frontal
and maxilla. The postorbital process of the frontal is large,
tapered, and extends posteroventrally to contact the end of
the zygomatic process of the squamosal. Within the posterior
part of the temporal fossa the surface of the parietal protrudes
laterally. The zygomatic process of the squamosal is large,
deep dorsoventrally, and approximately 55 mm long, mea-
suring from the suture with the exoccipital to the anterior
extremity. It does not diverge at an angle from the braincase
as in most living species of Delphininae, but, more like the
monodontids, has its long axis parallel to the midline of the
skull. Because the zygomatic process is set far laterally on
the cranium, there is a wide squamosal fossa between that
process and the lateral wall of the braincase. This squamosal
fossa forms a broad and concave floor of the temporal fossa
and has a thin and upturned anterior margin.

Distinctive characters of the zygomatic process of P. na-
salis are its narrow dorsal edge and prominent and square
posterolateral corner lateral to the paroccipital process. The
latter serves to buttress the glenoid fossa in the area posterior
to the nearly vertical postglenoid process. The fossa for the
middle air sinus, medial to the glenoid fossa, is partly un-
derhung by a thin, medial extension of the glenoid articular
surface for the mandible. A sharp, anteromedially directed
crest of bone separates this fossa from the cranial hiatus
(which held the ear bones). The homologous crest in most
other species of dolphins is developed into the elongate sty-
liform process that descends from the skull, but no indication
of such a process is present on any skull of P. nasa/is. A
small foramen (approximately 1 mm in diameter and which
can be probed with a needle to a depth of several mm) enters
the squamosal dorsal to the position of the external acoustic
meatus. The foramen is present on the paratype and holotype
of P. nasalis, and apparently is characteristic of this species,
although its homology and function are unknown. Posterior

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Barnes: Pithanodelphis from California 9

5cm

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10 Contributions in Science, Number 367

Barnes: Pithanodelphis from California

Figure 4. Pithanodelphis nasalis, new species, paratype skull, LACM 26635, LACM locality 5069; a, left lateral view; b, right lateral view.

5cm

to the glenoid fossa, on the ventral surface of the squamosal,
the location of the external acoustic meatus is marked by a
broad groove that is oriented transversely. A ventrally pro-
jecting part of the squamosal forms the posterior wall of this
groove, and is itself separated from the exoccipital by a fis-
sure.

Some ontogenetic changes can be noted within the avail-
able sample of skulls. Aside from obvious size increase these
include: relative enlargement of the nasal bones and corre-
sponding deepening of the depression between them ante-
riorly, fusion of the lateral margins of the premaxillae to the
maxillae in the area of the spiracular plates around the nares,
increase in prominence of the occipital and lambdoidal crests,
and increase in thickness of the lateral part of the maxilla
anterior to the antorbital notch.

Periotic. One of the two known periotics of P. nasalis is
the left one from the holotype (LACM 30093). The other
(LACM 122670) was found isolated. The cochlear portion
of the holotype periotic is crushed in its ventral and medial
sections, but the bone is otherwise intact. The periotic of P.
nasalis is notable by its compact appearance, i.e., the cochlear
portion is not prominent, the anterior and posterior processes
are small and do not project prominently from the bone, and
the whole periotic is somewhat flattened dorsoventrally. In
general proportions, absolute size, and relative positions of
structures, the periotics of P. nasalis somewhat resemble
those of an earlier, problematic fossil odontocete, Lampro-
lithax simulans Kellogg, 1931, a species that is known only
by isolated periotics from the Middle Miocene age Shark-
tooth Hill Bonebed in California. These two species show
differences of at least generic magnitude, however, and P.
nasalis has a relatively smaller cochlear portion with a small-
er internal acoustic meatus. In P. nasalis the cochlear portion
is not tilted so much anteriorly and does not have as prom-
inent a crease where it meets the medial surface of the anterior
process, the cerebral surface of the periotic lateral to the
cochlear portion is smoother and flatter, and the extremity
of the anterior process bends dorsally rather than ventrally.
Specimens of L. simulans (see Kellogg, 1931 :figs. 1 19, 120)
have a sinuous, elevated cerebral surface lateral to the coch-
lear portion, a more distinct and circular fossa for the head
of the malleus, and an articular facet for the tympanic bulla
which is three-sided, flattened posteriorly, and has an ex-
tremity which bends more laterally than posteriorly. In con-
trast, the periotic of P. nasalis has a four-sided posterior
articular surface (produced by acquisition of a comer on the
lateral edge) and the extremity of the posterior process points
posteriorly.

Unusual features of the P. nasalis periotic are an eminence
on the cochlear portion between the fenestra rotunda and the
cerebral orifice of aquaeductus cochleae, a prominent crest
on the posteromedial margin of the internal acoustic meatus,
the small size of the meatus, and the small, pointed anterior
process. The latter two characters, and the orientation of the
meatus, being twisted anteromedially, are similar to the peri-
otic of the holotype of the lophocetine kentriodontid Loph-
ocetus calvertensis ( Harlan, 1842) (see Barnes, 1978:fig. Ik),

Contributions in Science, Number 367

Barnes: Pithanodelphis from California 11

12 Contributions in Science, Number 367

Barnes: Pithanodelphis from California

Figure 6. Pithanodelphis nasalis, new species, holotype, skull, LACM 30093, LACM locality 5077, ventral view.

Contributions in Science, Number 367

Barnes: Pithanodelphis from California 13

Figure 7. Pithanodelphis nasalis, new species, paratype, skull, LACM 26635, LACM locality 5069, ventral view.

14 Contributions in Science, Number 367

Barnes: Pithanodelphis from California

Figure 8. Pithanodelphis nasalis, new species, composite reconstruction of skull based on available specimens, ventral view; for explanation of abbreviations see
Methods and Materials.

and might indicate some type of evolutionary relationship
between Pithanodelphis and Lophocetus Cope, 1867.

Tympanic bulla. Both tympanic bullae preserved with the
holotype are crushed and incomplete. The involucrum is
wide and massive compared with living dolphins in the gen-
era Delphinus Linnaeus, 1758, or Stenella Gray, 1866, and,
in its proportions, more resembles those of living porpoises
in the genera Phocoena Cuvier, 1817, or Phocoenoides An-
drews, 1911. On the ventral surface of the bulla the posterior
end of the involucrum is large and bulbous, the longitudinal
groove is broad and shallow, and a transverse constriction
separates the anterior and posterior parts. The anterior lip
of the bulla is rounded and there is no elongate styliform
process at the aperture of the auditory tube (eustachian tube
in part). The sigmoid process is bulbous and the posterior
process has an elongate projection posterior to the articular
facet for the periotic.

Ossicles. The malleus, incus, and stapes (Figs. 1 le-f) were
found in the matrix near the left periotic and bulla of the
holotype of Pithanodelphis nasalis. Ossicles are rarely found
and described for fossil cetaceans, but fortunately, the same
elements have been described for the holotype of Kentriodon
pernix (see Kellogg, 1927:28-31, hgs. 8-20), and the two
species may be compared. The anterior process of the mal-
leus, which in life was fused to the bulla, is incomplete (Fig.
1 1 f ), and might have been as large as in K. pernix. Compared
with K. pernix, the tubercle on the malleus of P. nasalis is
longer and the oblique groove cited by Kellogg is deeper.
This groove leads to a foramen ( fovea lateralis) for the chorda
tympani nerve that transects the bone from the juncture of
the two facets for the incus to the opposite (ventral) side.

The crus breve 'was, broken off of the incus. The crus longum
is shorter and wider than that of K. pernix, and the body of
the bone has wider articular facets for the malleus. The facet
for articulation with the stapes protrudes farther from the
side of the crus longum than in K. pernix.

The stapes of P. nasalis differs from that of K. pernix by
having a larger foramen (intercrural aperture), a wider head,
a more uniformly oval-shaped footplate, a smaller scar for
insertion of the stapedius muscle, and a more elongate facet
for the incus which is not set so obliquely on the bone.

The ossicles have been described for a more distantly re-
lated, but contemporaneous fossil delphinoid, the primitive
phocoenid, Salumiphocaena stocktoni (Wilson, 1973) (see
Barnes, 1977, 1984, 1985). That species has a malleus with
a wider anterior process and a shorter tubercle, and an incus
with a very reduced crus breve (Wilson, 1973:figs. 8a-d).
Based on comparisons of the ossicles of the three species,
Pithanodelphis nasalis, Kentriodon pernix, and Salumipho-
caena stocktoni, the former is the most primitive and the
latter is the most derived.

Mandible. Some oblique displacement of the right and left
dentaries shows that although the two sides were joined by
an extensive, rugose symphysis, they were not ankylosed.
The symphysis amounts to 40 percent of the length of the
mandible. The holotype bears alveoli for 22 teeth in the left
dentary, and 21 in the right, of which eight were posterior
to the symphysis in the left dentary and seven in the right.

Table 1. Measurements (in mm) of skulls of Pithanodelphis nas-
alis, new species. Parentheses indicate estimated measurements.

LACM

30093

Iloio-

type

LACM

26635

Para-

type

LACM

123872

Length of rostrum (2)

216

—

185

Width of rostrum at base (3)

78.5

87.5

73

Width of rostrum at midlength (5)
Width of premaxillae at midlength

28

—

26

of rostrum (6)

17

—

17

Greatest preorbital width (10)

(127)

(145)

(115)

Greatest postorbital width (11)

(145)

(156)

(115)

Least supraorbital width (12)
Greatest width of external nares

125

(138)

1 12

(13)

Greatest width across zygomatic

33

37.5

33

processes of squamosals (14)

(146)

166

—

Greatest width of premaxillae (15)
Greatest parietal width, within tem-

64.5

80

68

poral fossae (16)

79

99

—

Length of temporal fossa ( 1 9)

(85)

(82.5)

-

Width of temporal fossa (20)

(65)

68.5

—

Length of orbit (25)

Length of antorbital process of lac-

(45)

47

—

rimal (26)

20

21

15.5

Length of tooth row (32)
Number of teeth— left tooth row

188

—

168

(33)

Number of teeth — right tooth row

28

—

30

(34)

26

-

30

The tooth-bearing portion of each dentary is broad dorsally,
narrowly keeled ventrally, and bears two or three mental
foramina spaced along its lateral surface. The mandible is
slender and slightly up-turned anteriorly, and has a deeper,
somewhat keeled profile in the posterior symphyseal region.
Posterior to the alveolar rows the dentary expands dorsally
and ventrally and the bone in this part is thin and more
delicate. The posterior end of the coronoid process is directed
posteriorly and is separated by a concave mandibular notch
from the condyle. A slightly elevated coronoid crest 45 mm
posterior to the end of the alveolar row is turned slightly
laterally. The angle of the mandible extends farther poste-
riorly than does the coronoid process, but not farther than
the condyle. The condyle has a lateral buttress, is excavated
medially, and, when viewed posteriorly, has a vertical medial
margin and a convex lateral margin. On the medial surface
of the dentary, the anterior margin of the large mandibular
foramen extends to about the midlength of the post-sym-
physeal portion. The opening of this foramen (the mandib-
ular fossa) extends nearly from the dorsal to the ventral mar-
gin of the inner surface of the dentary.

Contributions in Science, Number 367

Barnes: Pithanodelphis from California 15

16 Contributions in Science, Number 367

Barnes: Pithanodelphis from California

Figure 10. Pithanodelphis nasalis, new species, paratype, LACM 26635, LACM locality 5069, posterior view, missing parts indicated by
dashed outline, natural size.

In the symphyseal portion of the mandible, the alveolar
rows are nearly parallel. They begin to diverge abruptly at
the posterior end of the symphysis. Interalveolar septa are
comprised of cancellous bone and are recessed between
prominent labial and lingual borders of the alveolar row.
Except for the posteriormost one or two, all of the alveoli
are about 4 or 5 mm in diameter and are directed dorsolat-
erally.

The mandible of Keniriodon pernix differs from that of P.
nasalis by being more slender and more elongate, having a
relatively shorter symphysis, a larger mandibular fossa and
nearly twice as many teeth. The only known mandible of
Deiphinodon dividum, that of the holotype, is incomplete,
missing its anterior end. Compared with P. nasalis, it appears
to have had a shorter symphysis and it has a higher, less

posteriorly projecting coronoid process, a larger mandibular
fossa, and more teeth in that part of each dentary which is
posterior to the symphysis.

Teeth. The dental formula on each side in P. nasalis is
26-30/21-22. The teeth in the middle parts of both the max-
illae and dentaries are approximately 1 5 mm to 20 mm long,
and the anterior and posterior ones are shorter. Each tooth
has a smooth, conical, enamel-covered crown that is curved
lingually at the apex. Teeth in the anterior and middle parts
of the alveolar row have nearly vertical crowns; the more
posterior ones have crowns that are shorter and more curved
lingually. All crowns bear a proximal lingual bulge that is
most prominent on the posterior teeth. The roots taper proxi-
mally, bend posteriorly and are bulbous below the gum line
(due to added outer layers of cement). No tooth in any of

Figure 9. Pithanodelphis nasalis, new species, referred specimen, skull, LACM 123872, LACM locality 6902; a, dorsal view; b. left lateral
view; c, ventral view.

Contributions in Science, Number 367

Barnes: Pithanodelphis from California 17

b d

Figure 11. Pithanodelphis nasalis, new species, left periotics and ossicles: holotype periotic, LACM 30093, LACM locality 5077; a, cerebral
or dorsal view; b, tympanic or ventral view; referred periotic, LACM 122670, LACM locality 6902; c, cerebral or dorsal view; d, tympanic
or ventral view; holotype ossicles, LACM 30093, LACM locality 5077; e, malleus, dorsomedial view; f, incus, ventral view; g, stapes, posterior
view; a-d, natural size, e-g, x5.

the skulls shows an open pulp cavity, indicating that the large
skulls are all of adults. There is no anteriorly directed, tusk-
like premaxillary tooth as in Kentriodon pernix (see Kellogg,
1927:pls. 2, 4, 5; Barnes and Mitchell, 1984:fig. 14a).

Vertebrae. The vertebral column of the paratype skeleton
(LACM 29087) is not complete, but includes the seventh
cervical, the first through sixth thoracic, the last three lumbar,
and 25 caudal vertebrae. Apparently there were at least one
and possibly two or three additional terminal caudal verte-
brae that were not preserved. An unknown number of tho-
racic and lumbar vertebrae were lost when the skeleton was
exposed by a trenching machine.

The spinous processes and the non-rib-bearing transverse
processes of these vertebrae are moderately elongate and flat.
They are not significantly expanded distally, as is the case in
some fossil and living odontocetes (see the unrelated, Recent
Pontoporia blainvillei (Gervais and d’Orbigny, 1844), for ex-
ample). The pedicles of the neural arches tilt anteriorly and
are positioned anteriorly on the centra. The pedicles of the
lumbar and anterior caudal vertebrae are thinner and flatter
than those of the thoracic vertebrae. The transverse processes
on the thoracic vertebrae are knob-like distally and have
shallow foveae for attachment of ribs. At least as far poste-
riorly in the body as the sixth thoracic vertebra, the capitulae
and tuberculae of the ribs were widely separated, judging by
the distance between their respective foveae on the vertebrae.
The transverse processes of the posterior lumbar and anterior
caudal vertebrae are positioned at mid-height on the centra.
Starting at the seventh caudal and continuing posteriorly,
each vertebra has a vertebrarterial canal on each side, located
posterior to the middle of the centrum. Caudals seven through
twelve retain spinous and transverse processes, and in each

of these vertebrae the canal pierces the transverse process
and is positioned ventral to the posterior edge of the pedicle
of the neural arch. After the twelfth caudal, the vertebrae
lack spinous and transverse processes, and become progres-
sively more rectangular in shape posteriorly. The eighteenth
caudal vertebra is significantly more expanded transversely
than the one immediately before it, and has rounded anterior
and posterior ends of its centrum. These features indicate
that this vertebra was the point of caudal flexion at the an-
terior margin of the caudal fluke. The vertebrae posterior to
this one are expanded transversely. Including the estimated
number of missing terminal caudal vertebrae, at least nine
vertebrae were originally included within the fluke.

Partial vertebral columns have been described for only two
other species of Kentriodontidae; the type specimens of Ken-
triodon pernix (see Kellogg, 1927) and of Delphinodon di-
vidum (see True, 1912b). The vertebrae of Pithanodelphis
nasalis differ from those of K. pernix by having spinous
processes that are higher and narrower and centra that are
more compressed anteroposteriorly and more expanded dor-
soventrally. The vertebrae of Delphinodon dividum are much
more like those of P. nasalis in both size and shape. Com-
pared with P. nasalis, the spinous processes of D. dividum
are slightly narrower anteroposteriorly and, in the anterior
thoracic region, these processes of D. dividum are more nearly
vertical and in the anterior caudal region they tilt more an-
teriorly. The centra of the anterior thoracic vertebrae of D.
dividum are slightly more compressed anteroposteriorly. Based
on these comparisons, the vertebral column of Pithanodel-
phis nasalis is more derived than that of Kentriodon pernix
and more primitive than that of Delphinodon dividum.

At least on the basis of vertebral structure, the living del-

18 Contributions in Science, Number 367

Barnes: Pithanodelphis from California

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Contributions in Science, Number 367

Barnes: Pithanodelphis from California 19

Figure 12. Pithanodelphis nasalis, new species, holotype, mandible, LACM 30093, LACM locality 5077; a, dorsal view; b, lateral view of left dentary.

Table 2. Measurements (in mm) of the holotype mandible, LACM
30093, of Pithanodelphis nasalis, new species.

Total length of dentary (38) (298)

Length of tooth row (37) (170)

Length of symphysis ( 1 20)

Height at coronoid process (39) 66

Length of mandibular fossa (40) 94

Number of teeth — left tooth row (35) 22

Number of teeth — right tooth row (36) 22

phinid species in the genera Sotalia and Sousa Gray, 1866,
are more primitive than Pithanodelphis nasalis in having
fewer vertebrae, longer centra, and wide, short spinous and
transverse processes. The same is true of otherwise relatively
derived living species of delphinids such as the pilot whales,
Globicephala spp.; false killer whales, Pseudorca crassidens
Owen, 1846; and killer whales, Orcinus orca Linnaeus, 1758.

In vertebral shape and proportions, a close living analog
of P. nasalis is the bottlenosed dolphin, Tursiops truncatus
Montagu, 1821. Although it is a larger animal, it is useful
for interpreting the vertebrae of the fossil. The paratype of
P. nasalis probably had 26 to 28 caudal vertebrae and in-
dividuals of T. truncatus have from 26 to 29 (Nishiwaki,
1963). Both species have approximately nine of these caudal
vertebrae within the flukes. In T. truncatus, the anteriormost
caudal vertebra that bears a vertebrarterial canal is the tenth
rather than the seventh as in P. nasalis. Caudals amount to
36 percent of the vertebral column in T. truncatus, and I
presume they comprised approximately the same percentage
in P. nasalis and that the remainder of the vertebrae in the
lumbar, thoracic, and cervical regions of the two species are
approximately proportional. Based on the measurements of
adult skeletons of T. truncatus, I calculated that the vertebral
column of the paratype of Pithanodelphis nasalis (LACM
29087) was approximately 155 cm long. Estimating that it
had a skull the length of that with the holotype (LACM
30093), and adding another cm for flesh covering the tip of
the rostrum, the probable total body length of the paratype
of P. nasalis in life was approximately 192 cm. As evidenced
by the large paratype skull, LACM 26635, some individuals
attained a larger size, and probably reached a total body
length of approximately 200 cm.

DISCUSSION

The Late Miocene species Phocaenopsis cornutus was first
briefly described by du Bus (1872:500), based on two frag-
mentary skulls from deposits in the Antwerp Basin, Belgium.
The genus to which he referred his species, however, Pho-
caenopsis Huxley, 1859, was originally based only on an
isolated humerus from New Zealand which Huxley thought
was Pleistocene in age. Fordyce (1981) has subsequently shown
that the type species of Phocaenopsis, P. mantelli, is Early
Miocene in age and that it belongs either in the family Rhab-
dosteidae (=Eurhinodelphidae) or Squalodontidae. du Bus
(1872:499) had also assigned another fossil species, Pho-

caenopsis scheynensis du Bus, 1872, to the same genus, but
both of these generic assignments were without sound basis
because of the non-comparable nature of the type materials.

Abel (1905:133) transferred Phocaenopsis scheynensis to
the genus Acrodelphis Abel, 1899, and (1905:142) Phocaen-
opsis cornutus to the new genus Pithanodelphis Abel, 1905.
Abel (1905:140-45, figs. 24-25) identified one of the two
skulls described by du Bus as the “original de Phocaenopsis
cornutus, du Bus.” This is the specimen that I now designate
as the lectotype of the species. Abel’s illustrations of the
dorsal and right lateral views of the skull and the accom-
panying text demonstrate the distinctive characters of the
species, du Bus had mentioned no other referred bones of
this species, but Abel (1 905:figs. 26-27) illustrated fused atlas
and axis vertebrae that he referred to the species, and in the
diagnosis (p. 143) stated that the atlas and axis were nearly
always fused, separate only in one example. In my opinion,
Abel’s referral of those cervical vertebrae to Pithanodelphis
cornutus is unfounded, because there are no demonstrated
associations with skulls.

The genus Pithanodelphis remained monotypic until the
present study, and no additional specimens have been re-
ferred to the type species. Abel originally classified it in the
subfamily Delphininae of the family Delphinidae. True
(19 12b: 192) retained Pithanodelphis cornutus in the family
Delphinidae and compared it with his new fossil species,
Delphinodon dividum. Winge (1921) did not contest the fa-
milial assignment of Pithanodelphis, but did observe that the
large medial extensions of the posterior ends of the maxillae
behind the cranial vertex was an unusual character when
compared with living species in the Delphinidae. (This con-
dition is characteristic of all kentriodontids.) Kellogg (1927)
named Kentriodon pernix as a new genus and species of del-
phinid and compared it with D. dividum. When Slijper ( 1 936)
named the Kentriodontinae as an extinct subfamily of the
Delphinidae, he included within it Kentriodon and Delphin-
odon and excluded Pithanodelphis. Simpson (1945) did not
recognize the Kentriodontinae, nor any other extinct or living
subfamilies of Delphinidae, but he did list Delphinodon, Ken-
triodon, and Pithanodelphis among the extinct genera of the
family. I recognized the Kentriodontidae as a separate family
(Barnes, 1978:25-26), and classified Pithanodelphis, Ken-
triodon, Delphinodon, and other genera in the nominate
subfamily, Kentriodontinae. As a result of the present study,
I now recognize substantial numbers of unique, derived fea-
tures of Pithanodelphis that warrant its classification within
a separate, new subfamily, the Pithanodelphinae.

I had previously characterized (Barnes, 1978) the family
Kentriodontidae, in part, as lacking asymmetry of the cranial
vertex, such as exists in all species of Delphinidae (sensu
stricto). Pithanodelphis nasalis and P. cornutus are, however,
kentriodontids that do, in fact, have asymmetrical cranial
vertices, but the extent of this asymmetry and the ways in
which the bones had become modified from the primitive
pattern are different from species in the Delphinidae. Among
the species of Delphinidae, the pattern of asymmetry and the
relationships of the bones that comprise the cranial vertex
are very uniform (Barnes, 1978:3), and the unique type of

20 Contributions in Science, Number 367

Barnes: Pithanodelphis from California

cranial asymmetry of species of Pithanodelphis differs in the
following ways. The posterior ends of the premaxillae are the
same length instead of the right one being longer. Both pre-
maxillae contact the nasals instead of only one. The posterior
end of the right premaxilla adjacent to the nares is only 2 to
4 mm wider than the left instead of being approximately
twice as wide. The nasal bones are high and peaked, forming
the highest part of the cranial vertex, not low and hemi-
spherical and forming the anterior side of the vertex. The
spiracular plates are of equal height, rather than the left being
more elevated than the right. The area of the cranial vertex
that is occupied by the right and left nasals is equal, rather
than the left nasal being smaller. The suture between the two
nasal bones twists to the right posteriorly, rather than to the
left. The mesethmoid septum between the nares lies on the
midline of the skull, instead of being offset to the left side.
The nares are equal in diameter, rather than the right being
larger.

In Pithanodelphis, as in species of Delphinidae, the pos-
terior end of the right maxilla extends farther toward the
midline than does the left, and the shapes of the right and
left halves of the occipital crest are different. Pithanodelphis
has additional unique, derived characters that differentiate
it from Delphinidae: the left nasal bone is lower than the
right, a very slender posterior extension of each premaxilla
is compressed between the maxilla and the nasal, and the
exposed area of the frontals behind the nasals has a shape
with five points.

The cranial asymmetry of Pithanodelphis is, therefore, of
a different nature than that which is found within the Del-
phinidae and such asymmetry was probably acquired inde-
pendently in the two groups from different ancestors that had
symmetrical cranial vertices. An as yet unnamed, contem-
poraneous species of true delphinid, which has an asym-
metrical cranial vertex of the type seen in living delphinids,
has been found in the same part of the Monterey Formation
as Pithanodelphis nasalis (see Barnes, 1977). The presence
of cranial asymmetry, as well as the well-developed spiracular
plates around the nares in Pithanodelphis, suggest the pres-
ence in life of some type of specialized musculature and nasal
passage diverticulae. In living odontocetes such structures
have been implicated in production of sound that is used in
echolocation (see Mead, 1975).

The retention of a fairly large olfactory fontanelle, as in
Pithanodelphis nasalis, is primitive and is a relatively un-
common occurrence among other species of fossil odonto-
cetes known after Middle Miocene time. Other primitive
cranial characters of the species are the large, laterally placed
zygomatic process of the squamosal, the long and tapered
postorbital process of the frontal, the exceptionally large tem-
poral fossa and the relatively small fossa for the pterygoid
sinus in the pterygoid hamulus. The relatively small size of
the paroccipital process is a derived character.

Pithanodelphis might have evolved from some taxon with-
in the subfamily Kentriodontinae because, in addition to the
family characteristics, it shares with the earlier Middle Mio-
cene species of Delphinodon and Kentriodon the following
characters: wide facial region, intermediate length rostrum

with premaxilla extending only a short distance anteriorly
beyond the maxilla, elongate postorbital process of the fron-
tal, transversely compressed and otherwise similarly shaped
zygomatic process of the squamosal, deep squamosal fossa
between the zygomatic process and the braincase, similar
distribution of air sinuses, and similar sizes and positions of
the basioccipital crests and cranial hiatus. Pithanodelphis
nasalis more specifically shares with D. dividum: a slightly
arched rostrum, similar tooth count, small paroccipital pro-
cess, convex lateral margin of the maxilla over the temporal
fossa, zygomatic process of the squamosal with tapered an-
terior end and projecting posterolateral comer, and similar
structure and proportions of vertebrae. Teeth without ac-
cessory denticles, and the unusually large nasal bones of P.
nasalis are derived characters compared with D. dividum.

Species in the genus Lophocetus also have relatively large
nasal bones, but these are shaped differently than those of
Pithanodelphis, being not significantly higher than the max-
illae, and more compressed transversely by them. The peri-
otics of both Lophocetus calvertensis (see Barnes, 1978:fig.
Ik) and Pithanodelphis nasalis share an oddly tapered an-
terior process, a wide lateral portion and a relatively large
internal acoustic meatus that is tilted anteriorly on the coch-
lear portion. It may be that the contemporaneous Late Mio-
cene species of Pithanodelphis and Lophocetus had closely
related Middle Miocene ancestors within the subfamily Ken-
triodontinae, such as the derived kentriodontine genus Del-
phinodon, and evolved in divergent ways.

Pithanodelphis nasalis and P. cornutus are the only species
assigned to the genus Pithanodelphis. Each species has a dif-
ferent combination of primitive and derived characters. My
interpretation of the polarity of these characters is based on
the anatomy of much more primitive Odontoceti such as
species in the families Agorophiidae, Squalodontidae and in
the kentriodontid subfamily Kampholophinae. The more de-
rived characters of P. cornutus include its reduced cranial
crests and more rounded, convex occipital shield. The de-
rived characters of P. nasalis include its larger nasal bones
and smaller, more tapered zygomatic processes of the squa-
mosals.

There are very few previously named fossil odontocetes of
Late Miocene age from the North Pacific realm (Barnes, 1977),
and most are known only by one specimen. In the existing
fossil cetacean collections of this age that have been obtained
from southern California deposits, however, Pithanodelphis
nasalis is the most abundant odontocete species and is now
represented by seven specimens that are mentioned in this
study. This is also the largest published sample of skulls of
any kentriodontid species. Such a relatively good sample
could, of course, be attributed to a collecting bias, but six of
the seven specimens came from a relatively small geographic
area in Orange County and from a restricted stratigraphic
interval in the Monterey Formation from which virtually all
noticeable vertebrate fossils, ranging from single bones to
complete skeletons, were collected and prepared. The only
specimen of P. nasalis recorded in this study that is not from
the Monterey Formation in Orange County is from the Mo-
delo Formation in the Santa Monica Mountains of Los Angeles

Contributions in Science, Number 367

Barnes: Pithanodelphis from California 21

Figure 13. Pithanodelphis nasalis, new species, paratype, LACM 29087, LACM locality 5082, cervical and thoracic vertebrae; cervical
vertebra 7 through thoracic vertebra 6; a, dorsal view; b, left lateral view; anterior views of individual vertebrae; c, first thoracic; d, third
thoracic; e, sixth thoracic. All to the same scale.

a— , i

5 cm

22 Contributions in Science, Number 367

Barnes: Pithanodelphis from California

Contributions in Science, Number 367

Barnes: Pithanodelphis from California 23

b, left lateral view.

County. This formation has been correlated with the Mon-
terey Formation in Orange County (Woodford et al., 1954:
fig. 2) and because both rock units were deposited in the
same marine basin, the Los Angeles Basin, all the specimens
may therefore be regarded as part of the same faunal aggre-
gate.

Furthermore, the relatively restricted source of the speci-
mens from the Monterey Formation reinforces the conclu-
sions based on morphological similarities that the specimens
attributed to P. nasalis represent one species. The sample of
skulls presents a range of size, but a consistency of mor-
phology. Within the sample of skulls of P. nasalis the vari-
ability (in the positions of the premaxillary and maxillary
foramina, in the size and shape of the nasals and the antor-
bital and postorbital processes, and in the size and anterior
extent of the fossae for the pterygoid sinuses) is commen-
surate with that in the small sample of three skulls of the
Atlantic kentriodontid, Kentriodon pernix, from the Calvert
Formation (Barnes and Mitchell, 1984).

Pithanodelphis nasalis was a contemporary of the earliest
known delphinid (sensu stricto; see Barnes, 1977:330), which
was recovered from the same part of the Monterey Formation
at Laguna Niguel. The earliest known phocoenid, Salumi-
phocaena stocktoni (Wilson, 1973), and two other kentri-
odontids, Lophocetus repenningi Barnes, 1978 and Liolithax
sp. (see Barnes, 1978), are also known from correlative rock
units in California.

Another contemporaneous odontocete, the small dolphin
Delphinavus newhalli Lull, 1914, was collected from the
Monterey Formation near Santa Maria, Santa Barbara Coun-
ty, California. It has been classified in the family Delphinidae
by Lull (1914) and Simpson (1945), but objectively consid-
ered, the present state of preservation and preparation of the
holotype and only known specimen preclude its assignment
to any particular family (Barnes, 1977 : 329). Delphinavus new-
halli is a much more derived species than Pithanodelphis
nasalis. In contrast with P. nasalis, it has 40 upper teeth
instead of 26 to 30, a more slender mandible with a sym-
physis approximately one-fourth as long, an edentulous an-
terior tip of the premaxilla that does not extend beyond the
end of the maxilla, and a shorter, thicker zygomatic process
of the squamosal that has a more dorsoventrally expanded
anterior end.

Hesperocetus californicus True, 1 9 1 2a, is another contem-
poraneous dolphin, and is even more problematic. It is known
only by a fragment of mandible that was found in the San
Pablo Formation in the San Francisco Bay area of California
(see Barnes, 1977). It differs from P. nasalis by being larger,
and by having a thicker mandible with much larger teeth
that have rugose, rather than smooth, enamel.

Several lines of evidence suggest that Pithanodelphis na-
salis was an offshore species that lived in deep water over
the continental shelf. Diatomaceous sediments are usually
considered to have been deposited in deeper waters over the
continental shelves (cf. Calvert, 1966), and all the known
fossil ocurrences of P. nasalis have been found either in such
sediments or in lenses of coarser clastic sediment that are
enclosed within diatomites. Furthermore, both the San Joa-

quin Hills and Santa Monica Mountains occurrences of P.
nasalis are at sites that, in Late Miocene time, were many
miles out in the marine basin from the inferred ancient shore-
lines (Woodford et al., 1954). Most of the specimens consist
of parts of associated or articulated skeletons or are otherwise
relatively unabraded, indicating that they were not trans-
ported great distances before their final deposition. Some
cetacean carcasses in the modem oceans have been known
to drift great distances prior to sinking to the sea floor (Schaf-
er, 1972:20-21) and it is possible, of course, that the Pith-
anodelphis nasalis fossils represent carcasses that were not
buried near the same environment in which they lived. It is
undoubtedly significant, however, that all the known speci-
mens have been found in the sedimentary context described
above.

Pithanodelphis nasalis had an overall adult body length of
approximately 200 cm. The vertebral column has propor-
tions very much like the living bottlenosed dolphin, Tursiops
truncatus. Its rostrum and teeth are also like that species and
other small living delphinids, and by analogy, its diet was
probably generalized, consisting mostly of small pelagic fish
and some squid (Rice, 1984:479). Such a diet would also be
consistent with the inference that P. nasalis was an offshore
species. Pithanodelphis nasalis was also a generalized animal
in its postcranial, mandibular, and dental morphology. Its
cranial asymmetry, moderate development of fossae for air
sinuses, and large mandibular fossa suggest that it was ca-
pable of echolocating. The orbits are relatively large com-
pared with Kentriodon pernix, indicating that P. nasalis had
large eyes. The teeth are large and deeply rooted. The largest
teeth are in the middle of the tooth row, and all teeth have
a considerable accretion of cementum on their roots. The
large temporal fossa indicates that there was strong temporal
musculature, which, in conjunction with the large teeth, long,
firmly ankylosed mandibular symphysis and large postgle-
noid process, suggests that P. nasalis could effect a strong
grasp with its jaws.

CLASSIFICATION

The classification presented below reflects the changes in
knowledge of the family Kentriodontidae subsequent to my
1978 publication and includes the taxa that were discussed
by Barnes and Mitchell (1984). Within genera, the more
primitive species are listed first, and in general, I follow this
arrangement throughout the classification. Certain aspects of
the anatomy of Leptodelphis stavropolitanus Kirpichnikov,
1954, Sarmatodelphis moldavicus Kirpichnikov, 1954, and
Microphocaena podolica Kudrin and Tatarinov, 1965, are
poorly known, and it is difficult to assign these genera to
established subfamilies. The new subfamily Pithanodelphi-
nae reflects the very derived characters of Pithanodelphis,
including the asymmetry of the cranial vertex, which is not
known among other species of Kentriodontidae and which
was acquired in a manner unlike that in other families within
the superfamily Delphinoidea. Oligodelphis azerbajdzanicus
was classified by Mchedlidze (1976) as a species of Delphi-
nidae, but appears to belong in the Kentriodontidae, and
should therefore be re-evaluated in detail.

24 Contributions in Science, Number 367

Barnes: Pithanodelphis from California

Class Mammalia Linnaeus, 1758
Order Cetacea Brisson, 1762

Suborder Odontoceti Flower, 1867

Superfamily Delphinoidea (Gray, 1821) Flower, 1 864
Family Kentriodontidae (Slijper, 1 936) Barnes, 1 978
Subfamily Kampholophinae Barnes, 1978
Kampholophos Rensberger, 1969
Kampholophos serrulus Rensberger, 1969.
Middle Miocene, California, U.S.A.
Liolithax Kellogg, 1931
Liolithax pappus (Kellogg, 1955) Bames,
1978. Middle Miocene, Maryland,
U.S.A.

Liolithax kernensis Kellogg, 1931. Middle
and Late Miocene, California, U.S.A.
Liolithax sp. Bames, 1978. Late Miocene,
California, U.S.A.

Subfamily Kentriodontinae Slijper, 1936
Kentriodon Kellogg, 1927
aff. Kentriodon. Bames and Mitchell, 1984.
Early Middle Miocene, California,
U.S.A.

Kentriodon pernix Kellogg, 1927. Middle
Miocene, Maryland, U.S.A.

Kentriodon obscurus (Kellogg, 1931) Bames
and Mitchell, 1984. Middle Miocene,
California, U.S.A.

Delphinodon Leidy, 1869 (in part)
aff. Delphinodon dividum True, 1912b. Bames
and Mitchell, 1984. Late Early and/or
Early Middle Miocene, Japan; Califor-
nia, U.S.A.

Delphinodon dividum True, 1912b. Middle
Miocene, Maryland and Virginia, U.S.A.
Subfamily Lophocetinae Barnes, 1978
Lophocetus C ope, 1868
Lophocetus repenningi Bames, 1978. Late
Miocene, California, U.S.A.

Lophocetus calvertensis (Harlan, 1 842) Cope,
1868. Late Miocene, Maryland, U.S.A.
Subfamily Pithanodelphinae, new subfamily
Pithanodelphis Abel, 1905
Pithanodelphis nasalis, new species. Late
Miocene, California, U.S.A.
Pithanodelphis cornutus (du Bus, 1872) Abel,
1905. Late Miocene, Belgium
Kentriodontidae, incertae sedis:

Oligodelphis Mchedlidze and Aslanova in
Mchedlidze, 1976

Oligodelphis azerbajdzanicus Mchedlidze and
Aslanova in Mchedlidze, 1976. Late
Oligocene, Azerbaidzhan S.S.R.,
U.S.S.R.

Sarmatodelphis Kirpichnikov, 1954
Sarmatodelphis moldavicus Kirpichnikov,
1954. Late Miocene, Moldavian S.S.R.,
U.S.S.R.

Leptodelphis Kirpichnikov, 1954

Leptodelphis stravropolitanus Kirpichnikov,
1954. Late Miocene, Stavropol, Russian
S.F.S.R., U.S.S.R.

Microphocaena Kudrin and Tatarinov, 1965
Microphocaena podolica Kudrin and Tatar-
inov, 1965. Late Miocene, Ukrainian
S.S.R., U.S.S.R.

CONCLUSIONS

Pithanodelphis nasalis is a new species of small fossil dolphin
classified in the new subfamily Pithanodelphinae of the ex-
tinct delphinoid family Kentriodontidae. The species is known
by fossil skulls and postcranial bones from the Monterey and
Modelo formations at about 33°30' and 34°05' north lati-
tudes, respectively, in the Los Angeles Basin in southern
California, U.S.A. Its abundance in collections suggests that
this dolphin was the most abundant odontocete cetacean
inhabiting the North Pacific Ocean off the coast of southern
California at approximately 10 to 11 million years ago. The
fossil material is sufficient to confidently differentiate P. na-
salis from previously named, contemporaneous small odon-
tocetes from California: Hesperocetus californicus True,
1912a; Delphinavus newhalb Lull, 1914; Salumiphocaena
stocktoni (Wilson, 1973); and Lophocetus repenningi Bames,
1978.

Pithanodelphis nasalis and P. cornutus (du Bus, 1872), an
approximately contemporaneous fossil species that was found
in the Antwerp Basin in Belgium, are the only species pres-
ently assigned to the genus and to the subfamily Pithano-
delphinae. Pithanodelphis might have evolved from some
taxon within the subfamily Kentriodontinae. Pithanodelphis
nasalis is similar to two well-known Middle Miocene Atlan-
tic kentriodontines, Kentriodon pernix Kellogg, 1927, and
Delphinodon dividum True, 1912b. Its vertebral structure is
more derived than that of the former, more primitive than
that of the latter, and its cranial structure is more derived
than both. Species of Pithanodelphis differ notably from all
other delphinoids by having extremely large, convex nasal
bones that comprise the highest part of the cranial vertex.

Pithanodelphis nasalis and P. cornutus are the only species
of Kentriodontidae that are known to have had cranial asym-
metry. The manner in which this asymmetry was expressed
is different, however, from species in the other families of
the superfamily Delphinoidea; the Monodontidae, Phocoe-
nidae, and Delphinidae; and this feature is, therefore, a con-
vergent derived character. Asymmetry was possibly acquired
separately in each of the modem delphinoid families (and in
other odontocete families as well).

The presence of cranial asymmetry and other derived char-
acters of Pithanodelphis nasalis, such as spiracular plates on
the premaxillae, moderate development of fossae for air si-
nuses in the basicranium, and large mandibular fossae, sug-
gest that the species could echolocate. It has a rostrum of
moderate length and a homodont dentition comprised of
conical-crowned teeth. Like most small Recent delphinids
with such features, it probably had a generalized diet con-
sisting mostly of small fishes and occasional squid. The mor-

Contributions in Science, Number 367

Barnes: Pithanodelphis from California 25

phology of its vertebral column is relatively primitive and,
in conformation and numbers of vertebrae, is approximately
analogous to that of the living bottlenosed dolphin, Tursiops
truncatus. Pithartodelphis nasalis was a smaller animal, how-
ever, with an overall body length at maturity of approxi-
mately 200 cm. The nature of the sedimentary deposits that
yielded the fossils and the preservation of the bones indicate
that the usual habitat of the species was probably offshore
in deep water over the continental shelf.

ACKNOWLEDGMENTS

I thank the North American Rockwell Land Corporation
(Narland), S.&S. Construction Company (a subsidiary of
Shapell Industries, Inc. of Beverly Hills, California), and Mrs.
Louise Hanson (formerly of Moulton Ranch) for permission
to collect fossils on their respective properties. The Natural
History Museum of Los Angeles County Foundation and the
Smithsonian Institution provided travel funds during the
course of this work. Fieldwork that yielded specimens de-
scribed in this study was sponsored by LACM, by its Foun-
dation, and by the National Geographic Society (Grant No.
2530-82). The Byron L. Crume Company of Corona, Cali-
fornia donated a tractor and operator for some excavation
work at the Laguna Niguel sites. Curation was supported by
the National Geographic Society and the National Science
Foundation (Grant No. BSR 82-18194). I thank William A.
Clemens, Jr., Donald E. Savage, J. Wyatt Durham, and Frank
C. Whitmore, Jr. for their advice on an earlier version of the
text which I originally prepared as part of a Ph.D. disserta-
tion. Robert L. Brownell, Jr., Samuel A. McLeod, and Ed-
ward Mitchell offered comments on the ideas expressed herein.
I thank Marion J. Bohreer, W. Earl Calhoun, Terry and Mi-
chael Pohl, Michael D. Quarles, and David P. Whistler for
collecting specimens used in this study. Bohreer and Calhoun
helped with locality data. Funds for much of the preparation
of specimens and drawing of illustrations were provided by
the University of California Museum of Paleontology, Berke-
ley, where Dorothy Dechant Boaz prepared fossils, and J.
Patricia Lufkin rendered the line drawings. John DeLeon of
the LACM prepared the photographs. Jo Anne Hankey and
Susan E. Barnes helped with manuscript preparation.

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tocene deposits of Calfomia. University of California
Publications in Geological Science 1 3(4):23— 1 32.

. 1925a. Additions to the Tertiary history of the

pelagic mammals on the Pacific coast of North America.

III. A new fossil sirenian from Santa Barbara County,
California. Carnegie Institution of Washington Publi-
cation 348(3):57-70, pi. 9, fig. 3, pis. 10-1 1.

. 1925b. Additions to the Tertiary history of the

pelagic mammals on the Pacific coast of North America.

IV. New pinnipeds from the Miocene diatomaceous earth
near Lompoc, California. Carnegie Institution of Wash-
ington Publication 348(4):71-96, pis. 12-13.

. 1927. Kentriodon pernix, a Miocene porpoise from

Maryland. Proceedings of the United States National
Museum 69( 1 9): 1—55, pis. 1-14.

. 1928. The history of whales— Their adaptation to

life in the water. Quarterly Review of Biology 3(1, 2):29-
76, 174-208.

. 1931. Pelagic mammals from the Temblor For-
mation of the Kern River region, California. Proceedings
of the California Academy of Sciences, Series 4, 19(12):
217-397.

Lawrence, B., and W.E. Schevill. 1956. The functional
anatomy of the delphinid nose. Bulletin of the Museum
of Comparative Zoology, Harvard College 1 1 4(4): 103—
51.

Lull, R.S. 1914. Fossil dolphin from California. American
Journal of Science, 4th Series 37(219):209-220, pi. 8.

Mchedlidze, G.A. 1976. Osnovnyye cherty paleobiologi-
cheskoy istorii kitoobraznykh. (Basic features of the pa-
leobiological history of cetaceans.) Akademia Nauk
Gruzinskoi S.S.R., Institut Paleobiologii, “Metsniereba”
Press, 136 pp., 32 pis. (In Russian, English summary.)

Mead, J.G. 1975. Anatomy of the external nasal passages

and facial complex in the Delphinidae (Mammalia: Ce-
tacea). Smithsonian Contributions to Zoology 207:i-iv,
1-72.

Miller, L.H. 1925. Avian remains from the Miocene of
Lompoc, California. Carnegie Institution of Washington
Publication 349:107-17, pis. 1-9.

Mitchell, E.D. 1968. The Mio-Pliocene pinniped Imago-
taria. Journal of the Fisheries Research Board of Can-
ada 25:1843-1900.

Nishiwaki, M. 1963. Taxonomical consideration on genera
of Delphinidae. Scientific Reports of the Whales Re-
search Institute, Tokyo 17:93-103.

Perrin, W.F. 1975. Variation of spotted and spinner por-
poise (genus Stenella) in the eastern Pacific and Hawaii.
Bulletin of the Scripps Institution of Oceanography,
University of California 2 1 : 1-206.

Repenning, C.A., and R.H. Tedford. 1977. Otarioid seals
of the Neogene. United States Geological Survey Profes-
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Rice, D.W. 1984. Cetaceans. In Orders and Families of
Recent Mammals of the World, eds. S. Anderson and
J.K. Jones, Jr., 447-90. New York: Wiley.

Schafer, W. 1972. Ecology and paleoecology of marine en-
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Simpson, G.G. 1945. The principles of classification and
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Slijper, E.J. 1936. Die Cetaceen. Vergleichend-anatomisch
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. 1912b. Description of a new fossil porpoise of the

genus Delphinodon from the Miocene formation of
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Philadelphia, 2nd Series 15:165-194, pis. 17-26.

Vedder, J.G., R.F. Yerkes, and J.E. Schoellhamer. 1957.
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Geological Survey, Oil and Gas Investigations, Map OM
193.

Wilson, L.E. 1973. A delphinid (Mammalia, Cetacea) from
the Miocene of Palos Verdes Hills, California. University
of California Publications in Geological Sciences 1 03: 1 —
34.

Winge, H. 1921. A review of the interrelationships of the
Cetacea. (Translated by Gerrit S. Miller, Jr.) Smithson-
ian Miscellaneous Collections 72(8): 1-97.

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Yerkes. 1954. Geology of the Los Angeles Basin. In
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Chapter 4. California Division of Mines Bulletin 170.

Accepted 1 April 1985.

Contributions in Science, Number 367

Barnes: Pithanodelphis from California 27

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Number 368

15 November 1985

EVOLUTIONARY RELATIONSHIPS OF THE
ATHERINOPSINAE (PISCES: ATHERINIDAE)

Brian N. White

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EVOLUTIONARY RELATIONSHIPS OF THE
ATHERINOPSINAE (PISCES: ATHERINIDAE)

Brian N. White1

ABSTRACT. The Atherinopsinae, a subfamily of the silverside fish
family Atherinidae, is cladistically diagnosed on the basis of five
synapomorphies: 1) Baudelot’s ligament ossified, 2) proximal end
of fourth ceratobranchial with moderate toothplates, 3) palatine with
medial bony shelf, 4) ventral projection of nasal bone contacting
lachrymal, and 5) lateral ethmoid with ventral bony knob. Two tribes
are recognized. The Atherinopsini is restricted to the Pacific shores
of North America and includes four genera: Atherinops, Athennopsis,
Colpichthys, and Leuresthes. Its members share three derived char-
acter states: 1 ) supraoccipital process trifid, 2) leading edge of vomer
with paired dorsal projections, and 3) expanded hypophyses formed
by bifurcation of haemal spines. The Basilichthyini is distributed
throughout the temperate marine and fresh waters of South America
and includes two genera: Basilichthys and Odontesthes. It is char-
acterized by three synapomorphies: 1) basioccipital fenestrated, 2)
extrascapular composed of two bony elements, and 3) haemal arches
expanded to form broad hypophyses. Within the northern tribe,
Leuresthes is most closely related to Atherinopsis whereas Atherinops
is considered to be most closely related to Colpichthys. The tropical
subfamily Menidiinae is considered to be the sister group of the
Atherinopsinae.

INTRODUCTION

After more than a century of effort, the evolutionary rela-
tionships of the silverside family Atherinidae have not been
adequately resolved. This study attempts to answer several
questions concerned with the evolutionary history of the Ath-
erinopsinae, an American subfamily. First, is the Atherinop-
sinae ( sensu Schultz, 1948) monophyletic? If so, what is its
sister group? Another question involves internal relation-
ships of the subfamily: is the Atherinopsinae composed of
phylogenetically distinct North and South American lin-
eages? The biogeographic history of the Atherinopsinae will
be considered in a separate contribution.

The New World atherinids have long been considered a
monophyletic group. The Atherinopsinae of Jordan and
Hubbs (1919) included almost all American silversides. In
a later revision, Schultz (1948:42) redefined the Atherinop-
sinae and restricted the subfamily “to that group of genera
now known from the Americas that have the premaxillary
dilated or broadened posteriorly and extending opposite or
into five or more of the broadened hypophyses of the haemal
arches, these specialized hypophyses mostly interconnecting

Contributions in Science, Number 368, pp. 1-20
Natural History Museum of Los Angeles County, 1985

with one another by flattish, broadened, spine-like bony pro-
cesses opposite the tapering part of the air bladder.” This
definition is based on a derived character having systematic
importance at the level of the New World atherinids and
excludes from the Atherinopsinae many genera included in
the subfamily by Jordan and Hubbs (1919), which Schultz
accommodated by the formation of a new subfamily, the
Menidiinae.

Only seven of the eight genera included in the Atherinop-
sinae by Schultz (1948) are still recognized: Atherinops (Stein-
dachner, 1876), Atherinopsis (Girard, 1854 ), Austromenidia
(Hubbs, 1918), Basilichthys (Girard, 1854), Colpichthys
(Hubbs, 1918), Leuresthes (Jordan and Gilbert, 1880), and
Odontesthes (Evermann and Kendall, 1906). Hubbsie/la
(Breder, 1936) was synonymized with Leuresthes by Moffatt
and Thomson (1975). The validity of some other atherinop-
sine genera, e.g. Colpichthys and Atherinopsis , have been
called into question (Todd, 1976).

Schultz (1948) did not consider the Menidiinae and Ath-
erinopsinae to be sister groups. Instead, he allied the Ath-
erinopsinae with the Old World subfamily Atherininae be-
cause a small number of atherinine species have haemal
modifications similar to those of some of the atherinopsine
fishes. He did admit, however, that this similarity may be
due to parallel evolution and have no bearing on the phy-
logenetic relationships of the two subfamilies. Jordan and
Hubbs (1919) proposed that the New World silversides (the
Atherinopsinae and Menidiinae of Schultz, 1948) and the
Old World Atherininae formed a monophyletic group.

Patten (1978) believed the Atherinopsinae of Jordan and
Hubbs (1919) to be monophyletic but did not consider this
assemblage to be related closely to the Atherininae. He also
questioned the division of the American atherinids into two
subfamilies, charging that Schultz (1948) used primitive
characters to define the Menidiinae.

The differing opinions of Patten (1978), Schultz (1948),
and Jordan and Hubbs (1919) raise questions about the evo-

1 . Research Associate, Section of Ichthyology, Natural History
Museum of Los Angeles County, 900 Exposition Blvd., Los Angeles,
California 90007.

ISSN 0459-8113

lutionary relationships of the American silversides which are
the basis for this study. Their resolution requires that the
New World silversides be viewed in a wider phylogenetic
context and that the Atherinopsinae be compared with out-
group species chosen from as many other teleostean taxa as
possible.

METHODS AND MATERIALS

The methods of Willi Hennig ( 1 966) are used herein to assess
evolutionary relationships. Known commonly as cladistics,
phylogenetic systematics, or cladism, this approach groups
organisms solely on the basis of shared derived character
states, or synapomorphies. Similarity due to the shared pos-
session of primitive or plesiomorphic character states is dis-
counted as uninformative and excluded from analysis.

Grouping organisms according to patterns of common an-
cestry demands that all taxonomic groups be monophyletic.
Put another way, all members of a taxonomic group must
share a common ancestor and all the descendants of that
ancestor must be included in the group. Synapomorphic char-
acter states are used to diagnose taxonomic groups. The hier-
archic arrangement of synapomorphic character states on a
cladogram serves to define the evolutionary relationships
between different groups.

Synapomorphic character states were determined by out-
group analysis (Lundberg, 1972; Watrous and Wheeler, 1981).
Outgroups were chosen primarily, but not exclusively, from
within the Atherinomorpha (Rosen and Parenti, 1981). In
outgroup analysis, character states restricted to the ingroup
are considered to be derived whereas states occurring in both
the ingroup and outgroup are assessed as being primitive.

In some cases, character states that are suspected of being
derived have limited occurrence among outgroup taxa and
must therefore be evaluated more critically. In such in-
stances, it is necessary to consider if it is more parsimonious
to assume that the character state in question was indepen-
dently derived in the ingroup and outgroup or whether it is
derived for a larger group and only retained by a small num-
ber of otherwise distantly related descendants. The assump-
tion that requires the fewest number of evolutionary steps,
the sum total of all gains and losses necessary to explain the
observed distribution of the character state, is accepted.

Arguments based on parsimony require a reasonable
knowledge of the overall relationship of the ingroup to related
lineages. For the purposes of this study, a recent phylogeny
proposed for the Atherinomorpha (White et al., 1984; Col-
lette, 1984) (Fig. 1) and the ideas of Patten (1978) concerning
the relationships of the Atherinidae (Fig. 2) were used to
settle questions of parsimony. In one case, the ontogeny of
a character was used to polarize a transformation series be-
tween three states.

At least two male and two female specimens of every ath-
erinopsine genus, except Basilichthys, were cleared and coun-
terstained (Dingerkus and Uhler, 1977) to facilitate obser-
vation of bone and cartilage. Only a single, male specimen
of Basilichthys was cleared and stained because of material
constraints. Osteological character states judged to be phy-

logenetically informative were examined further in a number
of partially dissected alcohol specimens. Material from nu-
merous outgroups was cleared and counterstained or dis-
sected as well.

The original description of every generic synonym is ref-
erenced in the synonymies presented in the discussion sec-
tion.

The preserved materials used in this study were furnished
by the following institutions: Natural History Museum of
Los Angeles County (LACM), Academy of Natural Sciences
of Philadelphia (ANSP), California Academy of Sciences
(CAS), National Museum of Natural History (USNM), Uni-
versity of Arizona (UA), University of California at Los An-
geles (UCLA), and University of Florida (UFj. The acronym
SU refers to collections of Stanford University now housed
at CAS. A list of species examined is provided below. Fol-
lowing each catalog number, in parentheses, are the total
number of specimens examined with the number of cleared
and stained preparations, if any, denoted by an asterisk.

Order Atheriniformes
Family Atherinidae
Subfamily Atherinopsinae

Atherinops affinis. LACM: 346(1), 347(1), 1808(1), 1809(1),
1984(1), 1995(1), 2619(1), 6609-1(1), 6612-2(1), 6615-2(1),
6616-1(1), 6635-2(1), 6683-1(1), 7990(1), 8823-8(1), 8909-
2(1), 8947-1 3(1), 9280-2(1), 928 1-1(1), 9297-1(1), 9380-2(1),
9439-2(1), 9592-3(1), 22075(1), 20125(1*), 22304(1),
22306(1), 30706-1(1), 31699-5(1), 32068-5(1), 32084-14(1),
32085-14(1), 32086-10(2,1*), 32184-14(9,1*), 32697-1(1),
32704-2(1), 33080-1(3), 33138-1(1), 33541-1(1), 351 53-10(1),
35794-1(1), 35815-1(1), 37013-1(4), 37552-5(1), 37575-
7(2,1*), 38545-2(1), 38548-1(1*), 82622-5(1), W48-34(l),
W49-157(l), W50- 144(1), W55-20(15), W55-90(l*), W56-
253(1), W63-59(l), W63-63(l), W68-43(21,2*).

Atherinopsis californiensis. LACM: 348(2,1*), 3896(1),
6735-2(1), 7936(3), 9283-1(1), 9439(3,1*), 20024(2),
20025(3,2*), 20120(1), 22300(1), 22302(1), 22795(1),
23227(1), 24066(1), 30636-6(1), 31306-2(6,3*), 31583-
4(6,1*), 31807-6(3), 31864-1(1), 31940-1(1), 32043-8(1),
32044-8(1), 32056-13(3), 32059-12(1), 32704-2(1*), 32925-
3(3), 32944-1(5), 33076-1(1), 34093-1(1), 37609-5(2), 42663-
4(2*), W49-4( 1 ), W49- 1 43(2), W58-377(42, 1 *), W67- 1 5 1 ( 1 ),
W67-152(l).

Basilichthys archaeus. USNM: 128536(2), 77530(1).

Basilichthys australis. LACM: 42705-1(1*); CAS:
SU22735(2); USNM: 84326(4).

Basilichthys semotilus. CAS: 45193(1), 45194(2),
SU23227(2).

Colpichthys regis. LACM: 7153(13,1*), 35728-1(10),
35730-1(10,4*), 39570-4(1*), W49-130(3), W50-190(2),
W51-15(10), W55-12(9,3*), W55-30(4).

Leuresthes sardina. LACM: 1523(1*), 9295(1*), 35728-
7(4), W49-121(l*), W50-18*(2,l*), W51-258(l).

Leuresthes tenuis. LACM: 1786(2), 1810(2), 4382(1),
4402(1), 6615(1), 6635-3(1), 6735-1(1), 8946-8(1), 9280-1(1),

2 Contributions in Science, Number 368

White: Systematics of Atherinopsinae

Figure 1. Relationships of the Atherinomorpha (White et al., 1984;
Collette, 1984).

/ \ \ \ /

/ \ \ \ /

\ \ \ V

y

Figure 2. Relationships of the subfamilies of the Atherinidae (Pat-
ten, 1978).

9453-1(1), 9592-2(1*), 20123(2), 20131(3,2*), 22307(2),
22879(1), 24070(2), 31306-3(2), 31757-7(1), 31759-22(2,1*),
32056-12(1), 32597-1(4,2*), 32946-1(1), 32947-1(2), 33077-
1(1), 33078-1(2), 33079-1(2), 33080-1(1*), 33139(1), 33487-
1(2), 37690-1(2), 38457-1(2,1*), W5 1-66(28,2*), W55-1 15(2),
W57-152(2), W66-62(3), W67-149A(1), W68-43(2), 015-SO-
08MA-01(4*), 024-RB-08-MA-0 1(4*), 024-SO-08-MA-
01(5*), 024-SO-22-MA-01(4*).

Odontesthes argentinense. USNM: 148502(4).

Odontesthes bonariensis. CAS: SU52812(4*).

Odontesthes brevianalis. USNM: 84338(1).

Odontesthes hatcheri. CAS: 12699(1), 42586(2).

Odontesthes incisus. CAS: SU31601(3); USNM: 163378(5).

Odontesthes mauleanum. CAS: 44702(2), 45201(2),
SU 1269 1(2); USNM: 77296(1), 84334(3).

Odontesthes nigricans. USNM: 77299(4), 88714(2),
103782(1).

Odontesthes perugiae. CAS: 11730(1).

Odontesthes regia. LACM: 20094(1), 42696-1(138,3*);
CAS: SU6072(1), SU9285(2), 1 1905(2), 45171(1), 45172(2),
45173(1), 45174(3); USNM: 77633(4), 77644(1).

Odontesthes smitti. USNM: 256719(10).

Undescribed species from Gulf of California. UCLA: W78-
11(4).

Subfamily Menidiinae

Archomenidia sallei. LACM: 43459-1(4).

Chirostoma grandocule. UCLA: W2-54(4).

Chirostoma labarcae. UA: 66-108-7(6).

Chirostoma sphyraena. UA: 66-128-1(2).

Coleotropis blackburni. LACM: 8335(2).

Eurvstole eriarcha. LACM: 1562(1*), 9044-16(4), 31784-
5(2*).

Hubbesia gilberti. LACM: 8964-2(1), 22328(7).

Labidesthes sicculus. LACM: 8965-1(2).

Melaniris chagresi. LACM: 9132-1(1*), 9148-9(1), 9167-
10(7).

Melanorhinus cyanellus. LACM: 20129(1), 35486-5(1*).

Membras martinica. LACM: 8975-1(1); ANSP: 125238(5);
UF: 35105(10).

Menidia beryllina. LACM: 8964-2(10).

Menidia peninsulae. LACM: 8962-2(2*).

Nectarges nepenthe. LACM: 20101(2*), 20103(1).
Poblana sp. LACM: 32616-1(4).

Xenatherina sp. LACM: 43458-1(4).

Xenomelaniris brasiliensis. ANSP: 120027(8).

Subfamily Notocheirinae

Iso rhothophilus. CAS: 46621(4).

Subfamily Atherioninae

Atherion elymus. LACM: W65-31(2).

Subfamily Melanotaeniinae

Pseudomugil signifer. LACM: 34988-3(2).

Subfamily Atherininae

Atherina breviceps. LACM: 42651-1(1*), 42695-1(4,1*).
Atherinomorus ogilbyi. LACM: 37481-1(13).
Atherinomorus pingnis. LACM: 31299-20(7).
Hypoatherina harringtonensis. LACM: 5833(2*).
Hypoatherina panatela. LACM: 42472-3(6).

Order Beloniformes
Family Belonidae

Pseudotylosurus angnsticeps. LACM: 41470-8(2).

Family Exocoetidae

Cypselurus opisthopus. LACM: 30455-1(3).

Exocoetus monocirrhus. LACM: 30473-5(4).

Contributions in Science, Number 368

White: Systematics of Atherinopsinae 3

Family Hemirhamphidae

Hyporhamphus unifasciatus. LACM: 6949-2(4).

Family Scomberesocidae

Cololabis saira. LACM: 34083-1(3).

Scomberesox saurus. LACM: 11223-1(4).

Strongylura timucu. LACM: 5875(8).

Order Cyprinodontiformes
Family Anablepidae

Anableps dowi. LACM: 42632-16(6*).

Oxyzygonectes dowi. LACM: 4876(3).

Family Cyprinodontidae

Belonesox belizanus. LACM: 42632-15(1).

Cyprinodon variegatus. LACM: 1309(5), 1310(4).

Floridichthys carpio. LACM: 1311(3).

Family Fundulidae

Fundulus diaphanus. LACM: 39823-1(4*).

Family Goodeidae

Goodea sp. LACM: 32615-1(2).

Xenotoca variata. LACM: 151(3).

Family Poecillidae

Poeci/ia sphenops complex. LACM: 9191-21(3).

Family Profundulidae

Profundulus guatemalensis. LACM: 1969-47(4).

Family Rivulidae

Rivulus isthmensis. LACM: 2779(3).

RESULTS

In all of the atherinopsine fishes examined, Baudelot’s liga-
ment is ossified at its point of attachment to the base of the
skull such that two thin, sharp spines are directed postero-
ventrally from the basioccipital (Fig. 3). As in most other
teleost fishes, the unossified portion of this ligament has a
point of attachment on the cleithrum. The placement, size,
and shape of these paired ossifications are constant through-
out the Atherinopsinae. Baudelot’s ligament is not ossified
in any of the other atheriniform species examined except Iso
rhothophilus. Species of the marine genus Iso are very deep
bodied and their overall appearance is quite different from
that of the Atherinopsinae. This dissimilarity carries over to
the ossification of Baudelot’s ligament, as the orientation of
the ossification on the base of the skull of I. rhothophilus
does not resemble that characteristic of the Atherinopsinae.

Figure 3. Ventral view of first two vertebrae and base of skull:
Colpichthys regis. OBL = ossified Baudelot’s ligament. Scale equals
1 mm.

In I. rhothophilus, the spines are placed higher on the skull
and are more laterally directed than in the atherinopsine
fishes. Therefore, the condition observed in I. rhothophilus
is considered to be independently derived and not homol-
ogous with the atherinopsine condition. Baudelot’s ligament
is ossified in each of the belonid, hemirhamphid, and scom-
beresocid species examined, but not in the exocoetids. In the
halfbeaks and sauries, the ossified ligaments form flat, sword-
like processes quite different from the cylindrical ossifications
typical of the Atherinopsinae. In the needlefishes, the ossified
portion of each ligament is much stouter than in the Ath-
erinopsinae. For these reasons, the beloniform pattern is not
considered to be homologous with the atherinopsine pattern.

A second derived character state shared by the atherinop-
sine fishes is found in the branchial basket: there are no
enlarged toothplates on the proximal end of the fourth cer-
atobranchial, but instead, there is a series of paired tooth-
plates running the length of the fourth ceratobranchial (Fig.
4a). In almost all atherinopsine species, these toothplates are
relatively narrow and do not contact their partners on the
dorsal midline of the fourth ceratobranchial. However, in
Colpichthys regis and in an undescribed atherinopsine col-
lected in the Gulf of California by Dr. Boyd Walker, these
paired toothplates are expanded so that they contact their
partners on the dorsal surface of the fourth ceratobranchial.
The typical atheriniform condition is seen in Menidia (Fig.
4b) in which an enlarged toothplate occurs on the proximal
end of the fourth ceratobranchial.

A third characteristic of the Atherinopsinae is found on
the palatine bone. In every atherinopsine species examined,
there is an ossified shelf on the medial side of the head of
the palatine bone. This shelf serves to brace the palatine
against the mesethmoid. It was not seen outside the Ather-
inopsinae. Nectarges and Atherinomorus (Fig. 5) were chosen
to illustrate the primitive atherinomorph condition of this
character. Ontogenetic stages of the palatine bone were ob-
served in cleared and stained larval specimens of Leuresthes
tenuis. The medial bony shelf appears after the head of the
palatine has developed the hammerlike shape shared by the
Atherinopsinae and Menidiinae.

4 Contributions in Science, Number 368

White: Systematics of Atherinopsinae

MBS

a

Figure 5. Posterior view of right palatine bone: a) Leuresthes te-
nuis, b) Nectarges nepenthe, c) Atherinomorus pingius. MBS = me-
dial bony shelf. Scale equals 1 mm.

A fourth derived feature of the Atherinopsinae concerns
the state of the nasal bone. The atherinopsine nasal bone has
a ventral projection (Fig. 6a) that makes contact with the
anterodorsal comer of the lachrymal bone. In other ather-
inomorphs, this projection is lacking and connection between
the nasal and lachrymal bones is accomplished via an ex-
tension of the anterior tip of the nasal sensory canal (Fig.
6b). In most atherinopsines, the ventral nasal projection is
well developed but in the California and Gulf grunions, re-
duction of the projection seems to have accompanied en-
largement of the lachrymal bone.

A fifth derived character state of the atherinopsine fishes
involves the lateral ethmoid. In the Atherinopsinae, there is
on the ventral surface of the lateral ethmoid a bony knob
(Fig. 7a) that abuts the posterior edge of the palatine. This
knob apparently provides additional support for the palatine
and presumably strengthens the entire snout. There is con-
siderable variation in this feature within the Atherinopsinae,
although it was well developed in all specimens examined.
It is most strongly developed in Atherinops affinis and least
developed in some specimens of Atherinopsis calif orniensis
(Fig. 7b). It was not encountered in any of the non-atheri-
nopsine species available for comparison.

Several synapomorphic osteological features occur in the
North American Atherinopsinae. The supraoccipital process
of most of the North American atherinopsines is trifid (Fig.
8a). In some specimens of Leuresthes tenuis the supraoccip-
ital process is bifid, but in L. sardina and the other North
American species the supraoccipital is strongly trifid. A trifid
supraoccipital process has been reported in several halfbeak
species (Collette, 1966). However, this contrasts with the
bifid condition of the supraoccipital process (Fig. 8b) of near-
ly all other atherinomorphs (Rosen, 1964). A bifid supra-
occipital process is considered here to be the primitive sil-
verside condition.

Figure 4. Dorsal view of fourth ceratobranchial dentition in the
Atherinidae: a) Atherinops affinis, b) Menidia peninsulae, c) Col-
pichthys regis. CB4 = fourth ceratobranchial. Scale equals 1 mm.

Contributions in Science, Number 368

White: Systematics of Atherinopsinae 5

Figure 6. Medial view of right nasal bones: a) Atherinopsis cali-
forniensis, b) Melanorhinus cyanellus. VP = ventral process. Scale
equals 1 mm.

Another derived character state shared by the North Amer-
ican genera occurs on the vomer. In the northern atherinop-
sines, a dorsal lip on the anterior edge of the vomer has a
pair of bony projections (Fig. 9) that contact the mesethmoid.
In other atherinids, there is either a single projection, as in
Menidia, or there is none at all, as in Iso (Patten, 1978).
Presumably, these projections help brace the mesethmoid.
The mesethmoid is a point of attachment for ligaments run-
ning to the palatine and maxillary bones and it is reasonable
to conclude that the additional support it receives from these
paired vomerine processes serves to strengthen the snout and
jaws of the atherinopsines of the northeastern Pacific.

One unique modification of the North American atheri-
nopsines involves the development of the haemal funnel into
which the swimbladder extends. The ontogeny of the broad,
haemal hypophyses begins with a cartilaginous haemal arch.
A groove develops on the ventral surface of the simple hae-
mal spines that deepens as ossification proceeds. The spines
finally bifurcate, with each half expanding to form a rect-
angular bony plate (Fig. 10a). Extending from the ventral
surface of each of these plates is a long, spinelike projection
that curves medially to meet its fellow from the opposite side
(Fig. 11a) (Schultz, 1948; Clothier, 1951). These spines do
not fuse, but form a second funnel ventral to the one formed
by the rectangular plates above them. It is into this second
funnel that the posterior end of the swimbladder projects.
This unusual modification of the anterior haemal arches leaves
them without a haemal spine and, in this way, the North
American atherinopsines differ from most other teleosts. A
similar condition is known to occur, however, in one other
atherinid, the atherinine species Atherinason hepsetoides
(Patten, 1978). Specimens of A. hepsetoides were not avail-
able for examination, but evidence presented by Patten (1978)
suggests that A. hepsetoides is distantly related to the North
American Atherinopsinae. Furthermore, none of the closest
relatives of A. hepsetoides has developed a similar condition.
I conclude that the haemal modifications of A. hepsetoides
and of the North American atherinopsines are independently
derived.

The form of the haemal arches is remarkably similar in
all of the northern atherinopsines except for the undescribed
species from the Gulf of California, which lacks these haemal
modifications. The body cavity of this species is truncate and
the swimbladder does not extend into the region of the caudal
vertebrae.

VK

b

Figure 7. Ventral view of right lateral ethmoid, anterior edge to
right: a) Atherinops affinis, b) Atherinopsis californiensis. VK = ven-
tral knob. Scale equals 1 mm.

The haemal modifications of the southern species are in
stark contrast with those of their northern counterparts. The
development of their hypophyses could not be observed be-
cause no larval material was available but the hypophyses

6 Contributions in Science, Number 368

White: Systematics of Atherinopsinae

a b

Figure 8. Dorsal view of supraoccipital process: a) Atherinopsis
californiensis, b) Menidia peninsulae. Scale equals 1 mm.

of the South American atherinopsines cannot be derived from
haemal spines, as they are in the North American atheri-
nopsines, because the haemal spines of the adult fishes are
intact. It appears that the haemal funnel occurring in the
southern species (Fig. 10b) develops from an expansion of
the haemal canals themselves instead of a bifurcation of the
haemal spines. There is only a single haemal funnel in the
southern atherinopsines, with the swimbladder extending into
the expanded lumens of a series of haemal arches that each
have a well-developed spine (Fig. 1 lb). The hypophyses of
the South American atherinopsines are furrowed delicately
and fenestrated in the same manner as the basioccipital bone
(Fig. 12). In some species, the hypophyses are quite complex,
as shown in Figure 10b, or they are simple. In Basilichthys
australis, for example, the hypophyses appear to be nothing
more than a simple flaring of the walls of the haemal canals.
A similar condition occurs in the atherinine species Atherina
boyeri, but it is not considered to be homologous with the
South American atherinopsine condition because other evi-
dence indicates that the Atherininae is distantly related to
the Atherinopsinae (Patten, 1978). In two South American
species, Odontesthes incisus and O. nigricans, the swimblad-
der is not posteriorly extended and the haemal arches are
unmodified.

Two other derived character states are shared by the South

PDF

a b

Figure 9. Dorsal view of vomer: a) Atherinopsis californiensis, b)
Atherinops affinis. PDP = paired dorsal processes. Scale equals 1
mm.

American atherinopsines. First, in all of the southern species
examined, the posterior portion of the basioccipital bone is
sculpted by a series of irregularly spaced foramina (Fig. 1 2).
Within the Atherinomorpha, this feature is found only in the
South American atherinopsines.

Another derived feature of the South American atheri-
nopsines involves the extrascapular bone. In most of the
southern atherinopsine species, the extrascapular is com-
posed of two bony elements (Fig. 13b, c), one oriented hor-
izontally and the other vertically. Both elements bear sensory
canals. In the North American atherinopsines, both sensory
canals are accommodated by a single bone (Fig. 1 3a). In most
atherinomorphs, the extrascapular bone is absent, but in some
genera, e.g. Menidia, it is fused with the posttemporal (Pat-
ten, 1978). Even though these two bones are fused in Me-
nidia, the direct communication between the two sensory
canals suggests that the single extrascapular bone of the North
American species is the ancestral condition for the subfamily.
This is not surprising, because in most fishes, the extrascap-
ular is composed of a single element (Weitzman, 1 962; Mead
and Bradbury, 1963; Springer, 1968; Zehren, 1979). There
is some variation in the form of the extrascapular bone in
the South American atherinopsines. In Basilichthys semo-
tilus, for example, the extrascapular bone resembles the North
American atherinopsine condition. This is not true of the
other species of Basilichthys examined. In three specimens,
two separate elements occurred. In one specimen, only one
element was present; the anterior vertical element was absent
on both sides. In another specimen having only one element,
the posterior horizontal elements were absent. In three other
specimens, both elements are present and united to form a
single bone. Because the development of two extrascapular
elements is widespread among the South American atheri-
nopsine genera, it is considered to be a derived feature uniting

Contributions in Science, Number 368

White: Systematics of Atherinopsinae 7

b

Figure 10. Lateral view of haemal hypophyses: a) Atherinops af-
finis, b) Odontesthes regia. Scale equals I mm.

them in a monophyletic group. The variation noted in Bas-
ilichthys semotilus and Odontesthes incisus is assumed to be
secondarily derived.

The menidiine fishes share several derived character states.
The parapophyses on the first vertebra are directed ante-
riorly, and are blunt distally (Fig. 14). The first parapophyses
of most other atherinomorphs are pointed and directed pos-
teriorly. In some cyprinodontiforms, notably Fundulus di-
aphanus, the parapophyses of the first few vertebrae project
forward and in others, such as Anab/eps dowi, the parapoph-
yses on most or all of the vertebrae are anteriorly directed.
In these cases though, the parapophyses are pointed and not
blunt as in the menidiines. Furthermore, only the menidiine
fishes have just the parapophyses on the first vertebra mod-
ified. There is variation in this feature within the Menidiinae.
In none of the menidiine genera examined, except Chiro-
stoma, are the parapophyses on the first vertebra pointed and
directed posteriorly, as in the atherinopsine fishes. Inspection
of juvenile specimens shows this to be a modification of the
typical menidiine pattern as the parapophyses on the first

b

Figure 11. Anterior view of caudal vertebra: a) Atherinops afftnis,
b) Odontesthes regia. HF = haemal funnel. Scale equals 1 mm.

vertebra of the smaller individuals are directed anteriorly
and are blunt.

In the Menidiinae and Atherinopsinae, the supraoccipital
canal extends over the frontal and pterotic bones. In both
subfamilies, three pores occur in the pterotic portion of the
canal. However, the two American subfamilies differ in the
number of pores that occur in the frontal portion of the canal.
The atherinopsine fishes have five frontal pores whereas the
menidiines usually have only four. When the supraorbital
pores are numbered according to the system of Gosline ( 1 949),
the atherinopsine total is seven (Fig. 1 5a) and the menidiine
total is six (Fig. 1 5b). There is some variation in the number
of supraorbital pores in the Menidiinae. The Mexican fresh-
water genus Poblana resembles the atherinopsine fishes in
having seven supraorbital pores. In Labidesthes sicculus, the
supraorbital canal lacks a bony roof so that no pores are
evident on the top of the head. This is true of the Old World
atherinid subfamilies as well. Parenti (1981) concluded that
it is primitive for the cyprinodontiform fishes to have seven
supraorbital pores and this is assumed to be the case in the
New World atherinids as well. There is great variation in the
number of supraorbital pores in the Beloniformes. In some
species, the cranial pores are miniscule and very numerous,
numbering up to 123 in the continuous supraorbital-post-
orbital-temporal canal of Scomberesox saurus (Parin and As-
takhov, 1982). In others, a pattern similar to the one seen in
the Atherinopsinae occurs. No beloniform species examined
during the course of this study was found to have the reduced
number of supraorbital pores characteristic of the Menidi-
inae.

Another derived menidiine character state involves the
enlarged toothplates present on the proximal end of the fourth
ceratobranchial (Fig. 4b). In all menidiine species examined,
these toothplates are fused to the ceratobranchial and cannot

8 Contributions in Science, Number 368

White: Systematics of Atherinopsinae

B

Figure 12. Lateral view of neurocranium: Odontesthes regia. B =
basioccipital. Open space blackened. Scale equals 1 mm.

Figure 13. Lateral view of right extrascapular bone: a) Atherinops
affinis, b) Odontesthes regia, c) Basilichthys australis. Scale equals 1
mm.

be dislodged from the branchial basket. In no other ather-
iniform for which information is available are the toothplates
attached in this manner (Patten, 1978). In the Cyprinodon-
tiformes, a series of teeth extends posteriorly along the dorsal
surface of the fourth ceratobranchial between the paired
toothplates that are arrayed along its right and left hand sides.
This does not resemble the menidiine pattern. Proximal
toothplates are absent in the Beloniformes.

Two unique osteological characters appear in both the Ath-
erinopsinae and Menidiinae. First, in both these subfamilies,
there is a modification of the pectoral girdle involving the
scapula and cleithrum. These two bones are connected in the
atherinopsine and menidiine fishes by a series of small, bony
buttresses. These buttresses resemble small foramina in lat-
eral view but are in fact bony struts passing between the
cleithrum and scapula. Some specimens have many struts
and others have few, but their shape, size and placement are
constant in the two subfamilies. The buttresses are always
elliptical in cross section and placed high along the dorso-
lateral surface of the cleithrum in a horizontal or nearly hor-
izontal arrangement. In Atherinomorus, the only other genus
in which bony struts were found to pass between the scapula
and cleithrum, the struts were vertically arranged along the
anterior edge of the cleithrum and are considered to be in-
dependently derived.

Another modification shared by the Atherinopsinae and
Menidiinae involves the palatine bone. In the fishes belong-
ing to these subfamilies, the head of the palatine is either
straight or shaped like a hammer. In Figure 16, the range of
variation exhibited by the palatine bone in New World ath-
erinids and the atherinomorphs as a whole is illustrated. In
Leuresthes (Fig. 16a), and all other atherinopsines, the head
of the palatine is shaped like a hammer. This is true of many
menidiines also, although in some, such as Nectarges (Fig.
16b), the palatine head is just bluntly rounded. During the
course of this study, the more typical atherinomorph con-
dition (Parenti, 1981), shown in the tapering, pointed pala-
tine of the Old World atherinid Atherinomorus (Fig. 1 6c) was
not encountered in any menidiine or atherinopsine species
examined.

The two grunion species share several derived character
states. First, neither Leuresthes tenuis nor L. sardina have
strongly developed teeth in the jaws as adults. It has been

reported that minute teeth do occur in these fishes (Moffatt
and Thomson, 1975) and that it is necessary to examine dried
skeletal material to observe them. No teeth were seen in any
of the cleared and stained or alcohol preserved specimens
examined in the course of this study. There is variation in
the dentition of the other atherinids. In Atherinopsis califor-
niensis, the jaw teeth are arranged in several rows, whereas
only a single row of teeth is found on the jaws of Atherinops
affinis. However, no other atherinid species are known to
have the weak dentition characteristic of the grunions. In
fact, most atherinomorphs have well-developed teeth in the
jaws.

A suite of derived traits is associated with the reproductive
biology of the grunions. Both Leuresthes tenuis and L. sar-
dina are lunar spawners that fertilize and bury their eggs at
the surf line on sandy beaches during the extreme high tides
of spring and early summer. The embryos develop in the
sand for approximately two weeks, when high tides again
reach the nests and initiate hatching (Walker, 1952). Though
a few other fishes are known to spawn on a lunar cycle, e.g.
Menidia menidia, the remarkable reproductive habits of the
grunions are a well-known specialization. The eggs of both
grunion species lack filaments. All other atherinopsine eggs
known have filaments as do the eggs of most atherinomorphs
(Rosen and Parenti, 1981; Collette, 1984).

The shape of the vomer in the California and Gulf grunions
is unusual (Fig. 17a). In both Leuresthes tenuis and L. sar-
dina, the leading edge of the vomer is emarginate and the
lateral condyles are reduced greatly. This contrasts with the
form of the vomer in the other atherinopsines and meni-
diines, where the lateral condyles are well developed and the

a b c d

Figure 14. Ventral view of anterior vertebrae and base of skull: a)
Menidia peninsulae, b) Nectarges nepenthe, c) Melaniris chagresi, d)
Chirostoma labarcae. Scale equals 1 mm.

Contributions in Science, Number 368

White: Systematics of Atherinopsinae 9

7

3

2

a

b

Figure 15. Supraorbital pore pattern: a) Atherinops affinis, b) Me-
lanorhinus cyanellus. Pores numbered according to the system of
Gosline (1949). Scale equals 1 mm.

leading edge has a strong median process (Fig. 1 7b). In overall
size, the vomer of the two grunions is reduced in comparison
with those of other atheriniforms.

The dorsal process on the premaxilla of L. tenuis and L.
sardina is long and slender and placed near the symphysis
of the upper jaw (Fig. 18a). The jaws are greatly protractile

Figure 16. Lateral view of right palatine: a) Leuresthes tenuis, b)
Nectarges nepenthe, c) Atherinomorus pinguis. Scale equals 1 mm.

in grunions and are not bound to the snout by a frenum as
in Basilichthys, Atherinops or Atherinopsis. A similar con-
dition is developed in Odontesthes (Fig. 18b), but in this
South American genus, the dorsal process is placed farther
back along the shaft of the premaxilla, suggesting that the
dorsal processes of the northern and southern genera were
independently derived.

Consideration of the other derived character states de-
scribed here suggests that the evolution of jaw mobility in
the Atherinopsinae involves the independent development
of slender, premaxillary dorsal processes and the loss of the
frenum to the upper jaw in the North American grunions
and the South American genus Odontesthes.

Atherinopsis californiensis shares with Leuresthes tenuis
and L. sardina another modification of the ethmoid region.
In these three species, the lateral ethmoid has a pointed, bony
strut that runs along the lateral edge of the parasphenoid (Fig.
19). This bony strut was not observed in any of the other
atherinid species examined. It is lacking in the beloniform
and cyprinodontiform fishes as well.

.LC

LC

MP

Figure 17. Ventral view of vomer: a) Leuresthes tenuis, b) Ather-
inops affinis. LC = lateral condyle, MP = median process. Scale equals
1 mm.

10 Contributions in Science, Number 368

White: Systematics of Atherinopsinae

DP

a

Figure 19. Ventral view of parasphenoid and lateral ethmoids:
Atherinopsis californiensis. P = parasphenoid, MP = median pro-
cess, LE = lateral ethmoid. Scale equals 1 mm.

b

Figure 18. Lateral view of right premaxillary bone: a) Leuresthes
tenuis, b) Odontesthes regia. DP = dorsal process. Scale equals 1
mm.

Another derived feature that Atherinopsis californiensis,
Leuresthes tenuis, and L. sardina have in common involves
the shape of the third epibranchial (Fig. 20a). The proximal
arm of the third epibranchial of these three fishes is twisted
about its long axis. Not only is the proximal arm of the third
epibranchial crooked, but it is noticeably thinner in Atheri-
nopsis californiensis, Leuresthes tenuis, and L. sardina than
it is in the other atherinopsines and menidiines, in which the
proximal arm of the third epibranchial is stout and uncon-
torted (Fig. 20b).

A single derived character state distinguishes Atherinopsis
californiensis from all other atherinomorphs. In A. califor-
niensis, there is a small pocket on the anterior tip of the
ventral process of the maxilla formed by a bony ledge pro-
jecting from its ventral surface (Fig. 23a).

Two unique osteological features are shared by Atherinops
affinis, Colpichthys regis, and the undescribed atherinopsine
species from the Gulf of California. In these fishes, the an-
terior edge of the quadrate is angled forward (Fig. 21b). This
feature is most apparent in dissected specimens because the
quadrate is held in position by the flesh that is digested away
in cleared and stained specimens. It can, of course, be ob-
served in cleared specimens, but mobility of the suspenso-
rium, of which the quadrate is a part, makes interpretation
of the exact orientation of the quadrate more difficult. In
none of the other atherinomorph genera examined was a
similar positioning of the quadrate observed. It is typical
within the Atherinomorpha for the anterior edge of the quad-
rate to have a vertical orientation (Fig. 21a).

The bicuspid teeth shared by Atherinops affinis, Col-
pichthys regis, and the undescribed species are unlike those
of any other atherinid (Schultz, 1948). In Atherinops affinis,

Figure 20. Left epibranchial: a) Atherinopsis californiensis, b) Ath-
erinops affinis. PA = proximal arm, UP = uncinate process. Scale
equals 1 mm.

Contributions in Science, Number 368

White: Systematics of Atherinopsinae 11

a

b

Figure 21. Orientation of the quadrate: a) Odontesthes regia, b)
Atherinops affinis. Q = quadrate. Cartilage stippled, open space
blackened. Scale equals 1 mm.

the cusps on each tooth are of equal size (Fig. 22b). In Col-
pichthys regis and the undescribed species, the cusps are un-
equal (Fig. 22a). Bicuspid and even tricuspid teeth are known
to occur in some cyprinodontiform fishes (Parenti, 1981) and
in the Hemirhamphidae as well.

Colpichthys regis and the undescribed atherinopsine from
the Gulf of California have in common several special fea-
tures involving the dorsal process of the maxilla and the
dentition of the branchial basket. There is a notch in the
dorsal process of the maxilla of both of these Gulf of Cali-
fornia atherinopsines (Fig. 23b). This notch was not seen in
any other atherinomorph species. Also shared by C. regis
and the undescribed species is the modification of the fourth
ceratobranchial toothplates described above. The toothplates
are wide and meet on the dorsal midline of the fourth cer-
atobranchial (Fig. 4c). Another unique feature seen in these
two species involves the teeth on the third pharyngobranchial
elements. These upper pharyngeal bones are covered with

a b

Figure 22. Jaw teeth: a) Colpichthys regis, b) Atherinops affinis.
Scale equals 0. 1 mm.

long hairlike teeth that apparently serve to strain fine sand
particles.

A single derived feature distinguishes Atherinops from all
other atherinomorph species: the fold of skin connecting the
distal ends of the premaxillary and maxillary bones is frilled
(Fig. 24).

Several unique character states occur in the South Amer-
ican atherinopsines. In the fishes assigned to the genus Basi-
lichthys, the ventral process of the maxilla has a rounded
projection on its dorsal surface that was not seen in any other
atherinomorph species examined (Fig. 23c).

Every species of Odontesthes or Austromenidia examined
lacks a mesethmoid. The Atherinomorpha is defined, in part,
on the presence of a disclike mesethmoid. Therefore, loss of
the mesethmoid is considered to be a derived character state
shared by these two genera.

Another derived feature shared by Odontesthes and Aus-
tromenidia involves the opercle. In every species examined,
except O. incisus, the opercle has on its anterodorsal comer
a complex array of foramina of varying sizes (Fig. 25). This
condition appears to be unique within the Atherinomorpha.
The absence of these foramina in O. incisus is judged to be
a secondary loss for two reasons. First, the occurrence of

12 Contributions in Science, Number 368

White: Systematics of Atherinopsinae

AP

a

Figure 23. Lateral view of right maxilla: a) Atherinopsis califor-
niensis, b) Colpichthys regis, c) Basilichthys australis. AP = anterior
pocket, DN = dorsal notch, RP = rounded process. Scale equals 1
mm.

FSF

Figure 24. Lateral view of head: Atherinops affinis. FSF = frilled
skin fold. Scale equals 1 mm.

these foramina is widespread among the other species in these
genera and second, the assumption that the foramina have
been lost in O. incisus is congruent with the other two derived
character states described above.

DISCUSSION

The synapomorphic character states described in the pre-
vious section support a phylogeny of the New World ath-
erinids (Fig. 26). Five derived character states (Node A) sup-
port the hypothesis that the Atherinopsinae ( sensu Schultz,
1948) is monophyletic: 1) Baudelot’s ligament ossified, 2)
proximal end of the fourth ceratobranchial without enlarged
toothplates, 3) palatine head with medial bony shelf, 4) ven-
tral projection of nasal bone contacting lachrymal, and 5)
lateral ethmoid with ventral bony knob.

The Menidiinae is proposed as the sister group of the Ath-
erinopsinae because they share seven synapomorphies: 1)
scapula and cleithrum connected by small bony struts, 2)
palatine head hammer shaped, 3) premaxilla with small an-
terior joint and broad alveolar arm, 4) premaxilla connected
to the coronoid by a short ligmanent 5) premaxilla free from
maxilla ventrally, 6) ethmomaxillary ligament attached to
palatine, and 7) posttemporal bone with flat anterior spine
extending into posttemporal fossa of skull (Patten, 1978).

Three derived character states support the contention that
the Menidiinae ( sensu Schultz, 1948) is monophyletic: 1)
parapophyses on the first vertebra directed anteriorly and
blunt distally, 2) number of supraorbital canal pores reduced
to four, and 3) toothplates fused to proximal end of fourth
ceratobranchial.

The internal relationships of the Atherinopsinae are clar-
ified by 18 derived character states. Three synapomorphies
characterize a South American assemblage (Node B): 1) ba-
sioccipital fenestrated, 2) extrascapular composed of two bony
elements, and 3) haemal arches expanded to form broad
hypophyses. A North American lineage (Node C) is defined
by three osteological modifications: 1 ) supraoccipital process
trifid, 2) a pair of dorsal projections on leading edge of vomer,
and 3) bifurcation of haemal spines forming expanded hy-
pophyses. Within the northern line one group composed of
the species of Atherinops and Colpichthys (Node D) is char-

Contributions in Science, Number 368

White: Systematics of Atherinopsinae 13

Figure 25. Medial view of right opercle: Odontesthes regia.
ADF = anterodorsal fenestration. Scale equals 1 mm.

acterized by two derived features: 1 ) leading edge of quadrate
angled anteriorly, not vertical, and 2) teeth bicuspid. Ath-
erinopsis and Leuresthes comprise a second lineage (Node E)
defined by two synapomorphies: 1) medial edge of lateral
ethmoid produced to form a spinelike projection contacting
lateral edge of parasphenoid, and 2) proximal arm of third
epibranchial contorted.

Colpichthys regis and the undescribed Gulf of California
species are proposed sister species sharing three derived char-
acter states: 1 ) dorsal process of maxilla notched, 2) teeth on
third pharyngobranchial long, hairlike, and 3) paired tooth-
plates on fourth ceratobranchial wide, gap between partners
much reduced. Atherinops can be diagnosed by the frilled
skin fold running between the distal ends of the maxilla and
premaxilla.

Two South American atherinopsine groups are recognized
on the basis of four synapomorphies. One assemblage, com-
prising the genus Basilichthys, is characterized by one osteo-
logical modification: ventral maxillary process with rounded
projection on dorsal surface. The second assemblage, which
includes all other South American atherinopsines, is defined
by three derived character states: 1) mesethmoid lacking, 2)
anterodorsal comer of opercle fenestrated, and 3) dorsal pro-
cess of premaxilla thin and spinelike, mouth protractile.

The evolutionary relationships illustrated in Figure 26 sug-
gest a change in the taxonomy of the Atherinopsinae. The
North and South American assemblages deserve tribal status
and names are proposed here for each group. The name
proposed for the northern tribe, the Atherinopsini, is taken
from Atherinopsis (Girard, 1854). The name proposed for
the southern tribe, the Basilichthyini, is taken from Basi-
lichthys (Girard, 1854). Both names are based on the first
genus described in the tribe. None of the North American
genera currently recognized need be synonymized to afford
sister groups equal taxonomic rank. Four northern genera

Figure 26. Generic relationships of the Atherinopsinae. Node A.
Baudelot’s ligament ossified, proximal end of fourth ceratobranchial
with moderate toothplates, palatine head with medial bony shelf,
ventral projection of nasal bone contacting lachrymal, lateral eth-
moid with ventral bony knob. Node B. Basioccipital fenestrated,
extrascapular composed of two bony elements, haemal arches ex-
panded to form broad hypophyses. Node C. Supraoccipital process
trifid, paired dorsal projections on leading edge of vomer, bifurcation
of haemal spines forming expanded hypophyses. Node D. Anterior
edge of quadrate angled forward, teeth bicuspid. Node E. Lateral
ethmoid with medial process contacting parasphenoid, proximal arm
of third epibranchial contorted. Node F. Maxilla ventral process
with rounded projection. Node G. Anterodorsal comer of opercle
fenestrated, mesethmoid lacking, mouth protractile. Node H. Skin
fold running from comer of mouth to maxilla frilled. Node I. Dorsal
process of maxilla notched, teeth on third pharyngobranchial hair-
like, toothplates on fourth ceratobranchial expanded. Node J. Ven-
tral process of maxilla with anterior pocket. Node K. Beach spawning
habits, mouth protractile, vomer reduced, dentition reduced or lack-
ing on jaws.

are recognized: Atherinops, Atherinopsis, Colpichthys, and
Leuresthes. Included in Colpichthys, is the undescribed species
from the Gulf of California. Two genera are recognized in
the Basilichthyini: Basilichthys and Odontesthes. Austro-
menidia is a junior synonym of Odontesthes.

Schultz ( 1 948) was correct in his conclusion that the Ath-
erinopsinae and Menidiinae are distinct evolutionary assem-
blages. The Atherinopsinae is diagnosed cladistically by five
characters; the Menidiinae by three. However, his contention
that the Atherinopsinae is most closely related to an Old
World subfamily, the Atherininae, is not supported by this
study.

The evidence presented here supports instead the hypoth-
esis that the Atherinopsinae and Menidiinae “represent a
lineage far removed from the ancestry of Atherininae” (Pat-
ten, 1978:99), do not form a monophyletic group, and con-
tribute two congruent, synapomorphic character states to the
definition of the New World line.

Apparently, similar osteological modifications have evolved
more than once in the New World atherinids. For instance,
in both the Basilichthyini and Atherinopsini, the swimblad-
der extends into a modified haemal funnel. Independent evo-
lution of this character is suggested by the fact that the mor-

14 Contributions in Science, Number 368

White: Systematics of Atherinopsinae

phology of the haemal funnel is quite different in the two
atherinopsine tribes. It is problematical whether the posterior
extension of the swimbladder in the northern and southern
tribes evolved independently. It could be that elongation of
the swimbladder occurred only once in the evolution of the
Atherinopsinae and that it is a derived character state shared
by all members of the subfamily. On the other hand, posterior
extension of the swimbladder and the haemal funnel in the
two tribes suggests that extension of the swimbladder and
modification of the haemal arches are correlated because
expanded haemal arches only occur in those species that have
the swimbladder elongated. It seems reasonable to assume
that if extension occurred only once in the Atherinopsinae,
then the haemal modification in the northern and southern
species would be identical. This argument is offered in sup-
port of the existence of separate North and South American
tribes within the Atherinopsinae, but since its truthfulness
cannot be objectively tested, posterior extension of the swim-
bladder is excluded from the diagnoses of the Atherinopsini
and Basilichthyini.

It is interesting to note that in the atherinid subfamily most
distantly related to the Atherinopsinae, the Atherininae, the
swimbladder extends into modified haemal arches in several
species. Some atherinine species, e.g. Atherina boyeri, ap-
proach the Basilichthyini in this regard. One species, Ath-
erinason hepsetoides, resembles quite closely the Atherinop-
sini. However, the haemal arches of the Notocheirinae,
Atherioninae, and Melanotaeniinae are unmodified and it
has been concluded that primitively the haemal arches are
unspecialized in the Atherininae (Patten, 1 9 7 8). It is apparent
that the similarities seen in the haemal arches of the Ath-
erinopsinae and some atherinine species were derived in-
dependently and are nonhomologous.

The premaxilla of the two North American grunion species
and the fishes of the South American genus Odontesthes seem
to have had independent origins as well. The premaxillae of
Basilichthys, Atherinops, Atherinopsis, and Colpichthys have
broadly based, triangular dorsal processes and the upper jaws
are not protrusile. The phylogenetic information discussed
above suggests that these are the most primitive taxa in the
Atherinopsinae and therefore, it is concluded that the con-
dition of their upper jaws is the ancestral state for the subfam-
ily. The upper jaws of Odontesthes, Leuresthes tenuis, and
L. sardina, on the other hand, are protrusile and the pre-
maxilla in these fishes has a long, slender dorsal process.
However, as has already been mentioned, the placement of
the elongate dorsal process in the northern and southern
species is different and jaw protrusion is considered on the
grounds of parsimony to have evolved more than once. With-
in the Atherinidae, there is a good deal of variation in the
form of the premaxillary dorsal process. The premaxillae of
the Atherinopsinae and Menidiinae are unique in that they
have only a small anterior joint (Patten, 1978). Therefore, it
would be unwise to compare jaw protrusion in the other
subfamilies with those of the New World atherinids because
the morphology of the upper jaws is different. However, it
is interesting that jaw protrusion has evolved several times
in the family (Patten, 1978) and that the genetic background

of these fishes allows for the development of analogous so-
lutions to the problem of jaw mobility.

Similar situations have been documented in other taxo-
nomic groups. In plethodontid salamanders, it has been dem-
onstrated that morphological specializations associated with
the tongue, limbs, and digits have evolved separately on
several occasions (Wake and Lynch, 1976; Larson et al.,
1981). The independent evolution of similar traits has also
been noted in the foot structure of gekkonine and diplodac-
tyline geckos (Russell, 1979) and in the adaptive ecology of
leptodactyline frogs (Martin, 1970).

Many systematists have studied parallel evolution and sev-
eral formal definitions have been proposed. According to
Simpson (1961:78), “Parallelism is the development of sim-
ilar characters separately in two or more lineages of common
ancestry and on the basis of, or channeled by, characteristics
of that ancestry.” Mayr (1969:202) defined parallelisms as
“similarities resulting from joint possession of independently
acquired phenotypic characteristics produced by a shared
genotype inherited from a common ancestor.” Hecht and
Edwards (1976:654) stated that in parallel evolution “the
character is present in the ancestral form but a common
derived state has been independently evolved in each de-
scendant form.” Nelson (1978:123) proposed that “parallel-
ism can be defined as the presence in two monophyletic taxa
of a common character state which has been derived through
identical successive character state changes from a dissimilar
state present in the most recent common ancestor of both
taxa.”

The independent development of an association between
the swimbladder and the haemal funnel in the Basilichthyini
and Atherinopsini fits all of these definitions but the last. To
meet the requirements of Nelson’s definition, the haemal
modifications of the Basilichthyini and Atherinopsini would
have to develop in exactly the same manner and be identical
in all respects. However, if this were true, it would be difficult
to recognize that the northern and southern condition were
independently derived. In fact, they would probably be viewed
as a single derived character state shared by both tribes,
definitive for the subfamily, but uninformative about its in-
ternal relationships.

It has been suggested that “the concept of parallelism be
omitted from systematic studies” and “the term convergence
be applied to all cases of nonhomologous character similar-
ities” (Eldredge and Cracraft, 1980:74). In the case of the
atherinopsine fishes, the haemal modifications of the Basi-
lichthyini and Atherinopsini clearly are nonhomologous, but
the development of a haemal funnel is a striking similarity
that presumably evolved from the primitive condition held
by their immediate common ancestor. I believe it misleading
to call this an example of convergent evolution because the
haemal morphology of the two tribes is divergent. However,
the Atherinopsini and Basilichthyini evolved comparable
haemal specializations that have the same function from a
genotype inherited from their most recent common ancestor.
This developmental potential passed on by the ancestral ath-
erinopsine was expressed differently in its two descendant
lineages, but they evolved in similar directions nonetheless,

Contributions in Science, Number 368

White: Systematics of Atherinopsinae 15

probably because of some unknown genetic or epigenetic
constraints. In this sense, parallel evolution has occurred in
the Atherinopsinae and, questions of process aside, I find the
definitions of Simpson (1961) and Mayr (1969) are mean-
ingful and useful.

KEY TO THE GENERA OF
ATHERINOPSINE FISHES

la. Supraoccipital process trifid (Fig. 8a) (sometimes bifid

in Leuresthes tenuis)', vomer with paired dorsal projec-
tions (Fig. 9); air bladder extending into haemal funnel
composed of modified haemal spines (Figs. 10a, 11a);
extrascapular composed of single bony element (Fig. 1 3a);
basioccipital not fenestrated. Atherinopsini 2

lb. Supraoccipital process bifid (Fig. 8b); vomer without

paired dorsal projections; air bladder extending into hae-
mal funnel composed of modified haemal arches (Fig.
1 Ob, lib); extrascapular composed of two bony elements
(Fig. 13b, c) (a single element in Basilichthys semotilus
and some Odontesthes incisus)', basioccipital fenestrated
(Fig. 12). Basilichthyini 5

2a. Jaw teeth bicuspid (Fig. 22); leading edge of quadrate
angled anteriorly (Fig. 2 1 b); proximal arm of third epi-
branchial straight not contorted (Fig. 20b); lateral eth-
moid without medial process contacting parasphenoid

3

2b. Jaw teeth unicuspid; leading edge of quadrate vertical
(Fig. 21a); proximal arm of third epibranchial crooked,
contorted (Fig. 20a); lateral ethmoid with medial process

contacting parasphenoid (Fig. 19) 4

3a. Ventral process of premaxillary with bony pocket on
anterior tip (Fig. 23a); vomer not reduced; jaws not pro-
tractile; jaw teeth not reduced Atherinopsis

3b. Ventral process of premaxillary without bony pocket on
anterior tip; vomer reduced; jaws greatly protractile; jaw

teeth minute or absent Leuresthes

4a. Tooth cusps equal in length (Fig. 22b); skin fold between
comer of mouth and maxilla frilled (Fig. 24); dorsal
process of maxilla without notch; toothplates on fourth
ceratobranchial not expanded (Fig. 4a) .... Atherinops
4b. Tooth cusps unequal in length (Fig. 22a); skin fold be-
tween comer of mouth and maxilla not frilled; dorsal
process of maxilla with notch (Fig. 23b); toothplates on

fourth ceratobranchial expanded (Fig. 4c)

Colpichthys

5a. Mesethmoid absent, mouth protractile; anterodorsal
comer of opercle fenestrated (Fig. 25) (except in Odon-
testhes incisus)', ventral process of maxilla without
rounded projection on dorsal surface .... Odontesthes
5b. Mesethmoid present; mouth not protractile; anterodor-
sal comer of opercle not fenestrated; ventral process of
maxilla with rounded projection on dorsal surface (Fig.
23c) Basilichthys

CLASSIFICATION AND SYSTEMATIC ACCOUNTS

Subfamily Atherinopsinae

Tribe Atherinopsini
Genus Atherinops Steindachner, 1876
Genus Atherinopsis Girard, 1854
Genus Colpichthys Hubbs, 1918

Genus Leuresthes
Jordan and Gilbert, 1880

Tribe Basilichthyini

Genus Odontesthes

Evermann and Kendall, 1906

Genus Basilichthys Girard, 1854

Subfamily Atherinopsinae Fowler, 1904

DIAGNOSIS. Baudelot’s ligament ossified; proximal end
of fourth ceratobranchial with moderate toothplates; palatine
head with medial bony shelf; ventral projection of nasal bone
contacting lachrymal; lateral ethmoid with ventral bony knob.

COMPOSITION. Six genera: Atherinops Steindachner,
1876; Atherinopsis Girard, 1 854; Basilichthys Girard, 1854;
Colpichthys Hubbs, 1918; Leuresthes Jordan and Gilbert,
1880; Odontesthes Evermann and Kendall, 1906.

DISTRIBUTION. West coast of North America from
Vancouver Island to the Gulf of California in marine waters.
Temperate South America in fresh and marine waters.

Tribe Atherinopsini Fowler

DIAGNOSIS. Supraoccipital process trifid; vomer with
paired dorsal projections; haemal funnel developed from bi-
furcated haemal spines.

COMPOSITION. Four genera: Atherinops Steindachner,
1 876; Atherinopsis Girard, 1854; Colpichthys Hubbs, 1918;
Leuresthes Jordan and Gilbert, 1880.

DISTRIBUTION. Marine waters of western North Amer-
ica from Vancouver Island into the Gulf of California.

Genus Atherinops Steindachner

Atherinops Steindachner, 1876:89 (type species Atherinopsis

affinis Ayres, by monotypy).

DIAGNOSIS. Atherinopsines with fold of skin running
from comer of mouth to distal end of maxilla frilled.

DESCRIPTION. Baudelot’s ligament ossified; proximal
end of fourth ceratobranchial with moderate toothplates; pal-
atine head with medial bony shelf; ventral projection of nasal
bone contacting lachrymal; lateral ethmoid with ventral bony
knob; supraoccipital process trifid; vomer with paired dorsal
projections; haemal funnel comprised of modified haemal
spines; jaw teeth bicuspid; anterior edge of quadrate angled
forward.

16 Contributions in Science, Number 368

White: Systematics of Atherinopsinae

COMPOSITION. One species, the type.
DISTRIBUTION. Outer coast and islands of Oregon, Cal-
ifornia, and Baja California.

Genus Atherinopsis Girard

Atherinopsis Girard, 1 854: 1 34 (type species Atherinopsis cal-
iforniensis Girard, by monotypy).

DIAGNOSIS. Atherinopsines with pocket on anterior tip
of ventral process of maxilla.

DESCRIPTION. Baudelot’s ligament ossified; proximal
end of fourth ceratobranchial with moderate toothplates; pal-
atine head with medial bony shelf; ventral projection of nasal
bone contacting lachrymal; lateral ethmoid with ventral bony
knob; supraoccipital process trifid; vomer with paired dorsal
projections; haemal funnel comprised of modified haemal
spines; proximal arm of the third epibranchial contorted;
lateral ethmoid with medial process contacting parasphe-
noid.

COMPOSITION. One species, the type.
DISTRIBUTION. Outer coasts and islands of North
America from Vancouver Island to Baja California.

Genus Colpichthys Hubbs

Colpichthys Hubbs, 1918:67 (type species Atherinops regis
Jenkins and Evermann, by original designation).

DIAGNOSIS. Atherinopsines with notch in dorsal process
of maxillary bone; toothplates on fourth ceratobranchial ex-
panded; teeth on third pharyngobranchial hairlike.

DESCRIPTION. Baudelot’s ligament ossified; proximal
end of fourth ceratobranchial with moderate toothplates; pal-
atine head with medial bony shelf; ventral projection of nasal
bone contacting lachrymal; lateral ethmoid with ventral bony
knob; supraoccipital process trifid; vomer with paired dorsal
projections; haemal funnel composed of modified haemal
spines; jaw teeth bicuspid; anterior edge of quadrate angled
anteriorly.

COMPOSITION. Two species: C. regis and a new species
from Gulf of California.

DISTRIBUTION. Gulf of California.

Genus Leuresthes Jordan and Gilbert

Leuresthes Jordan and Gilbert, 1880:29 (type species Ath-
erinopsis tenuis Ayres, by monotypy).

Hubbsiella Breder, 1936:6, figs. 2-4 (type species Menidia
clara Evermann and Jenkins, by monotypy).

DIAGNOSIS. Atherinopsines with beach spawning hab-
its; jaw teeth reduced or absent; mouth greatly protractile;
vomer reduced.

DESCRIPTION. Baudelot’s ligament ossified; proximal
end of fourth ceratobranchial with moderate toothplates; pal-
atine head with medial bony shelf; ventral projection of nasal
bone contacting lachrymal; lateral ethmoid with ventral bony
knob; supraoccipital process trifid; vomer with paired dorsal
projections; haemal funnel composed of modified haemal

spines; proximal arm of third epibranchial contorted; lateral
ethmoid with medial process contacting parasphenoid.
COMPOSITION. Two species; L. sardina and L. tenuis.
DISTRIBUTION. Outer coasts and islands of California,
Baja California, and the Gulf of California.

Tribe Basilichthyini, New

DIAGNOSIS. Basioccipital ridged and fenestrated; ex-
trascapular composed of two bony elements; haemal funnel
developed from expanded haemal canals.

COMPOSITION. Two genera: Basilichthys Girard, 1854;
Odontesthes Evermann and Kendall, 1906.

DISTRIBUTION. Temperate South America in marine
and fresh waters including high Andean streams.

Genus Basilichthys Girard

Basilichthys Girard, 1854:198 (type species Atherina micro-
lepidota Jenyns, by subsequent designation of J ordan, 1919).
Protistius Cope, 1874:66 (type species Protistius semotilus
Cope, by monotypy).

Gastropterus Cope, 1878:700 (type species Gastropterus ar-
chaeus Cope, by monotypy).

Pisciregia Abbott, 1899:342 (type species Pisciregia beards-
leei Abbott, by monotypy).

DIAGNOSIS. Atherinopsines with rounded projection on
ventral process of maxilla.

DESCRIPTION. Baudelot’s ligament ossified; proximal
end of fourth ceratobranchial with moderate toothplates; pal-
atine head with medial bony shelf; ventral projection of nasal
bone contacting lachrymal; lateral ethmoid with ventral bony
knob; basioccipital ridged and fenestrated; extrascapular
composed of two bony elements; haemal funnel developed
from expanded haemal canals.

COMPOSITION. Approximately 6 species; generic re-
vision needed.

DISTRIBUTION. Marine and fresh waters of Peru and
Chile.

Genus Odontesthes Evermann and Kendall

Odontesthes Evermann and Kendall, 1906:94, fig. 3 (type
species Odontesthes perugiae Evermann and Kendall, by
subsequent designation of Jordan and Hubbs, 1919).
Kronia Ribeiro, 1915:9 (type species Kronia iguapensis Ri-
beiro, by monotypy).

Pseudothyrina Ribeiro, 1915:1 1 (type species Pseudothyrina
iheringi Ribeiro, by monotypy).

Austromenidia Hubbs, 1918:307 (type species Basilichthys
regillus Abbott, by original designation).

Cauque Eigenmann, 1928:56 (type species Chirostoma mau-
leanus Steindachner, by original designation).

Patagonina Eigenmann, 1928:56, 60 (type species Patagonia
hatched Eigenmann, by monotypy).

Patagonia Eigenmann, 1928:56 (l.c. in footnote, lapsus pro
Patagonina, takes same type).

Contributions in Science, Number 368

White: Systematics of Atherinopsinae 17

? Austroatherina Marrero, 1950, not seen, after de Buen 1 953
(type species Atherina incisa Jenyns).

Yaci de Buen, 1953:51, figs. 31, 32, 33 (type species Yaci
retropinnis de Buen, by monotypy).

DIAGNOSIS. Atherinopsines with mesethmoid lacking,
mouth protractile, anterodorsal comer of opercle fenestrated.

DESCRIPTION. Baudelot’s ligament ossified; proximal
end of fourth ceratobranchial with moderate toothplates; pal-
atine head with medial bony shelf; ventral projection of nasal
bone contacting lachrymal; lateral ethmoid with ventral bony
knob; basioccipital ridged and fenestrated; extrascapular
composed of two bony elements; haemal funnel developed
from expanded haemal canals.

COMPOSITION. Approximately 10 species; generic re-
vision needed.

DISTRIBUTION. Temperate South America in marine
and fresh waters from Peru to southern Brazil. Also Malvinas
and Juan Fernandez Islands.

ACKNOWLEDGMENTS

I thank Robert J. Lavenberg for his guidance and generous
support of this research. I hope this manuscript in some small
way repays my enormous debt. Camm Swift was an inex-
haustable source of information on the anatomy, osteology,
and relationships of fishes. Barry Chemoff generously pro-
vided unpublished phylogenetic information. Walter Ivan-
tsoff and John Patten made available their unpublished thes-
es which served as the foundation for my work. Without
their cooperation, my task would have been much more
arduous. Robert Lea and the late John Fitch provided study
material and shared their opinions on the relationships of
the silversides. Donald Buth advised me on phylogenetic
techniques and kindly loaned some obscure and out of print
references. My ideas were influenced greatly by conversations
with fellow graduate students: Martin Meisler, Gerald
McGowen, Steven Caddell, Richard Feeney, William Szel-
istowski, Noelle Sedor, Michael Miyamoto, Steven Werman,
Dana Krempels, Marc Hayes, Maureen Donnelly, Craig
Guyer, and C. Ben Crabtree.

The following people provided gifts or loans of specimens:
Jeffrey Seigel (Natural History Museum of Los Angeles
County), William Smith-Vaniz (Academy of Natural Sci-
ences of Philadelphia), Tomio Iwamoto (California Academy
of Sciences), Susan Jewett (National Museum of Natural His-
tory), Philip Hastings (University of Arizona), Donald Buth
(University of California, Los Angeles), Carter Gilbert (Uni-
versity of Florida), and Philip Heemstra (J.L.B. Smith In-
stitute, Grahamstown, South Africa).

Any errors in fact or interpretation are my own and in no
way reflect the views of those who so generously aided and
encouraged this research.

This study was partially funded by Biomedical Research
Support Grant 5 S07 RR070 12, Division of Research Re-
sources, Bureau of Health Professions, Education and Man-
power Training, National Institutes of Health.

LITERATURE CITED

Abbott J.F. 1899. The marine fishes of Peru. Proceedings
of the Philadelphia Academy of Sciences 51:324-364.

Ayres, W.O. 1 860. (original description of Atherinopsis af-
finis). Proceedings of the California Academy of Sciences
2:73-86.

Breder, C.M. 1936. Scientific results of the second ocean-
ographic expedition of the “Pawnee” 1926. Bulletin of
the Bingham Oceanographic Collection 2:6-10.

Buen, F. de. 1953. Los pejerreyes (familia Atherinidae) en
la fauna Uruguaya, con descripcion de nuevas especies.
Boletim do Instituto Oceanografico 4(l-2):3-80.

Clothier, C.R. 1951. A key to some southern California
fishes based on vertebral characters. Fish Bulletin, no.
79, 83 pp. California Division of Fish and Game.

Collette, B.B. 1966. Belonion, a new genus of fresh-water
needlefishes from South America. American Museum
Novitates 2274, 22 pp.

. 1984. Atherinomorpha: Introduction. Page 334 in

Ontogeny and Systematics of Fishes, ed. H.G. Moser,
W.J. Richards, D.M. Cohen, M.P. Fahay, A.W. Kendall,
and S.L. Richardson. Special Publication no. 1, 760 pp.
American Society of Ichthyologists and Herpetologists.

Cope, E.D. 1874. Description of some species of reptiles
obtained by Dr. John F. Bransford, assistant surgeon
United States Navy, while attached to the Nicaraguan
surveying expedition in 1873. Proceedings of the Acad-
emy of Natural Sciences of Philadelphia 26:64-109.

. 1878. Synopsis of the fishes of the Peruvian Am-
azon, obtained by Professor Orton during his expedi-
tions of 1873 and 1877. Proceedings of the American
Philosophical Society 17:673-701.

Dingerkus, G., and L.D. Uhler. 1977. Enzyme clearing of
alcian blue stained whole small vertebrates for dem-
onstration of cartilage. Stain Technology 52:229-232.

Eldredge, N., and J. Cracraft. 1980. Phylogenetic Patterns
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Evermann, B.W., and O.P. Jenkins. 1891. Report upon a
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with descriptions of new species. Proceedings of the
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Evermann, B.W., and W.C. Kendall. 1906. Notes on a
collection of fishes from Argentina, South America, with
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Fowler, H.W. 1904. Descriptions of new, little known, and
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Girard, C. 1854. Descriptions of new fishes collected by
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18 Contributions in Science, Number 368

White: Systematics of Atherinopsinae

Gosline, W.A. 1949. The sensory canals of the head in some
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Jordan, D.S. 1919. The genera of fishes, part II, from Agas-
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Contributions in Science, Number 368

White: Systematics of Atherinopsinae 19

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Accepted 24 April 1985.

20 Contributions in Science, Number 368

White: Systematics of Atherinopsinae

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A REVIEW OF THE GENERA OF CLEPTOPARASITIC
BEES OF THE TRIBE ERICROCINI
(HYMENOPTERA: ANTHOPHORIDAE)

Roy R. Snelling' and Robert W. Brooks1 2

ABSTRACT. The genera comprising the New World bee tribe Er-
icrocini are reviewed. Each genus is separated by a key, is described,
its included species-group names listed, and pertinent morphological
features illustrated. Nine genera are recognized: Mesoplia ( =Melissa )
and its new subgenus Eumelissa (type species, Melissa decorata F.
Smith), Hopliphora (=Eurytis = Oxynedys = Cyphomelissa), Me-
sonychium ( =Epiclopus ), Ericrocis, Abromelissa (new genus; type
species, Melissa lendliana Friese), Aglaomelissa (new genus; type
species, Melissa duckei Friese), Ctenioschelus (=Melissoda), Meso-
cheira, and Acanthopus.

A cladistic analysis of the Ericrocini is included.

Known hosts are listed in a table.

INTRODUCTION

The Ericrocini are a New World tribe of cleptoparasitic bees
largely centered in the Amazonian Basin. So far as known,
all hosts are within the related tribe Centridini. Despite their
attractive appearance, these moderate- to large-sized bees
have received scant attention from taxonomists, probably
due largely to their rarity in collections.

HISTORICAL RESUME

The first ericrocine genus to be described was Acanthopus,
proposed by Klug ( 1 807) for a single South American species.
Lepeletier and Serville (1825) described Mesonychium
(monobasic) and Mesocheira (three species). Ctenioschelus
was described by Romand (1840) for a single, bizarre species
and in 1841 Lepeletier described Me/issoda (later shown to
be isogenotypic with Ctenioschelus through synonymy), Me-
soplia and Hopliphora. Shuckard (1840) named Ischnocera,
with no included species, but it has long been recognized as
an obvious synonym of Ctenioschelus.

F. Smith (1854) added two new genera, Eurytis (mono-
basic), and Melissa (four species). All these genera were placed
in his subfamily Denudatae, together with such genera as
Melecta, Tha/estria, and Liogaster. He recognized Eurytis,
Melissa ( =Mesop/ia ), Mesocheira, Ctenioschelus ( =Melis -

Contributions in Science, Number 369, pp. 1-34
Natural History Museum of Los Angeles County, 1985

soda = Ischnocera) and Acanthopus. The one Nearctic genus,
Ericrocis, was described by Cresson ( 1 887) for a single species.

Ashmead (1899) assigned these bees to the family No-
madidae, recognizing the genera Ericrocis, Eurytis (=Hopli-
phora ), Melissa, Mesocheira, Mesonychium, and Acanthopus
( =Ctenioschelus ). A few years later, Schrottky (1902) pro-
duced his key to the Brazilian genera of Nomadidae, rec-
ognizing Melissa, Eurytis, Mesocheira, Ctenioschelus, and
Acanthopus, to which he added two new genera: Cyphome-
lissa and Oxynedes.

For the next 40 years, Schrottky’s generic concepts were
more or less followed by Cockerell, Ducke, and Friese, the
principal describers of Neotropical bees during that time.
Generic limits were flexible and considerable confusion ex-
isted regarding the application of the names Melissa, Me-
sonychium, and Mesoplia. These three names were very in-
consistently used, but in general Melissa was used to contain
most species in preference to Mesoplia, and Mesonychium
fell heir to those species which seemed not to be Mesoplia.

The Nearctic genus, Ericrocis, was included in the Melec-
tinae by Linsley (1939) who noted, however, its uniqueness
and commented that it is “perhaps ... an offshoot from some
group like Epicharis, which Grutte considers to be ancestral
to Acanthopus and Rathymus .” As Linsley noted, Cockerell
and Atkins (1902) had earlier emphasized the unusual fea-
tures of Ericrocis and related genera, and suggested that a
separate subfamily might be appropriate for these bees.

When Michener (1944) reorganized the higher classifica-
tion of the bees, he placed Ericrocis, and such similar genera
as Acanthopus, Mesocheira, and Ctenioschelus, in the tribe
Ericrocini, near the Centridini. He clearly set forth the dif-

1 . Entomology Section, Natural History Museum of Los Angeles
County, 900 Exposition Blvd., Los Angeles, California 90007, U.S.A.

2. Department of Entomology, University of Kansas, Lawrence,
Kansas 66045, U.S.A.

ISSN 0459-81 13

ferences between the Ericrocini and such cleptoparasitoid
groups as Melectini, Epeolini, and Nomadini.

The last author to deal with the group, more or less as a
whole, was Moure ( 1 946). The geographic scope of this work
was limited to Brazil, but since nearly all the genera occur
there, the treatment was nearly complete. Moure recognized,
and separated in a key, Ctenioschelus, Mesonychium, Me-
soplia, Hopliphora, Cyphomelissa, Acanthopus, and Oxyne-
dis (a misspelling of Oxynedys). Under Moure’s concepts,
Mesoplia included Melissa, and Epiclopus fell into Meso-
nychium.

TERMINOLOGY

In general, the morphological terminology follows that es-
tablished by Michener (1941), Michener and Fraser (1978),
and Winston (1979). A few terms used here necessitate ex-
planation. The interantennal distance is the shortest distance
between the inner margins of the antennal sockets. The an-
tennal socket diameter is the maximum transverse distance
across the antennal socket, from inner margin to outer mar-
gin. The antennocular distance is the shortest distance be-
tween the outer margin of the antennal socket and the inner
eye margin.

As in most bees, the short anterior face of the mesepi-
stemum and the much longer lateral face meet in a curved
surface. Sometimes, this juncture is marked by a raised Ca-
rina, the anterior mesepisternal carina (amc. Fig. 70). In most
ericrocines that possess this carina, it is thin, translucent, and
lamelliform. Ventrally, the anterior mesepisternal carina usu-
ally is confluent with a carina that bounds the posterior por-
tion of the procoxal cavity; this is the acetabular carina (ace,
Fig. 70) (Bohart and Menke, 1976). In a few ericrocines (e.g.,
Acanthopus), there is a low, rounded, shiny ridge which be-
gins in front of the metacoxa and extends for a short distance
onto the lateral face of the mesepistemum, the sternopleural
ridge (spr. Fig. 70). In most genera, the mesepistemum slopes
abruptly toward the coxal cavity, but there is no definite shiny
ridge. The supraspiracular ridge originates dorsolaterally on
the propodeum and extends posteriorly, above the propodeal
spiracle. The ridge is said to be strong if it terminates in a
blunt tooth or projection, weak if it becomes evanescent
apicad.

Male genital structures are somewhat confusing. The gono-
stylus is, in dorsal view, short, broad and more or less flat-
tened. In some genera the gonostylus is simply a very broad,
somewhat flattened structure, bearing diagnostic gonostylar
setae, but with little, if any, dorsal lobe. Dorsal lobes are
present in such genera as Mesoplia, Abromelissa, and Acan-
thopus; they may be present or absent in Hopliphora. When
present, the dorsal lobe is usually narrow, thin, lightly scler-
otized and always setose (Fig. 22). A second, much shorter
and broader, lobe may also be present immediately above
the base of the longer lobe.

In some genera there is a distinctive, heavily sclerotized
plate along the inner, basal portion of the gonostylus. No
similar structure is known within the Centridini or Rha-

thymini. For want of a better term this structure is here called
the inner apical sclerotization of the gonocoxite.

SYSTEMATICS

Although there are parasitic bee species in other families, the
greatest diversity, in numbers of genera and of species, is
within the Anthophoridae. One subfamily, the Nomadinae,
is exclusively parasitic and includes the majority of the species
in such large genera as Nomada, Hypochrotaenia, and Epeo-
lus. The few remaining parasitoid groups are mostly in the
Anthophorinae: Melectini, Rhathymini, and Ericrocini.

The most conspicuous difference between these three tribes,
as a group, and the worldwide Nomadinae is that females of
Nomadinae, with some exceptions (especially in the Old
World), possess a distinct, usually beveled, prepygidial fim-
bria or brush on the distal portion of the fifth abdominal
tergum. Most Nomadinae females, and often the males as
well, have a conspicuous, sharply defined, pygidial plate that
is commonly about one-half as broad at the base as the width
of the sixth tergum. Females of the Nomadinae (except Hex-
epeolus ) have only five exposed metasomal sterna (six ex-
posed sterna in cleptoparasitic Anthophorinae) and the legs
are commonly spiculate or tuberculate. In both sexes the
second abscissa of vein M+Cu of the hind wings is usually
at least twice as long as the (usually) transverse cu-v; in those
groups in which the second abscissa is not twice as long as
cu-v, the labrum is conspicuously longer than broad. The
apical portions of the wings are not papillate, as they are in
the Ericrocini.

The Melectini are a worldwide group that includes the
genera Melecta and Thyreus and a few smaller genera. Me-
lecta is a Holarctic genus that appears to be limited to north-
ern temperate regions and Thyreus is an exclusively Old
World genus that is primarily southern, reaching South Af-
rica and Australia. The hosts are mostly within the related
pollen-gathering tribe Anthophorini, from which the Melec-
tini are presumed to be derived.

Characteristics by which the Melectini differ from the Er-
icrocini are: the marginal cell barely, or not at all, exceeds
the last submarginal cell; the mesotibial spur is not modified;
the male gonostyli are slender and elongate; the mesobasi-
tarsus is more or less rounded in cross section and is without
a cariniform ridge along the posterior margin, whereas, in
the Ericrocini they are laterally flattened, with a cariniform
ridge (except Acanthopus and Hopliphora ); the labrum is
about as long as broad (except Zacosmia) and has a distinct
basal bulla on either side; and, the meso- and metatibiae of
the females are provided with coarse, spine-like setae.

The Rhathymini are exclusively Neotropical; there are fewer
than half a dozen species, all placed in the genus Rhathymus.
Known hosts are species of Centridini. The Rhathymini are,
like the Ericrocini, presumed to be derived from the Cen-
tridini and the two groups have many shared character states,
e.g., the general pattern of the wing venation, the lack of
spine-like setae on the meso- and metatibiae of the females,
the configuration of the scutellum and of the face, as well as
other features. Rhathymini differ from Ericrocini in the pres-

2 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

ence of basal bullae on the labrum, the wings are hairy, the
papillae are smaller and have hairs, the mesotibia! spur is
unmodified, and the body is without appressed, metallic scale-
like hairs.

Tribe Ericrocini

Ericrocini Cockerell and Atkins, 1902:46. Michener, 1944:

288; 1954:145.

Ctenioschelini Michener, 1965:10. NEW SYNONYMY.

Medium-sized to large, robust bees, often with distinct
patterns of white and iridescent blue, green or purple reflec-
tive scales and/or hairs, especially on abdomen.

Head conspicuously narrower than thorax, which is broad-
er than long or deep; labrum broader than long, with trans-
verse preapical ridge or median tubercle; clypeus about twice
wider than long, apical margin broadly concave; lateral angle
confluent with inner eye margin or nearly so. Malar space
virtually absent. Mandible simple or with a single preapical
tooth; posterior angle below middle of lower end of eye.
Postflabellum present. Maxillary palpus with 1-4 segments.
Antenna short, except in male Ctenioschelus; scape robust,
shorter than combined lengths of first three flagellar seg-
ments; first flagellar segment short, little, if any, longer than
broad (except male Ctenioschelus).

Pronotum short, collar closely appressed to front of meso-
scutum; scuteilum usually bituberculate. Three submarginal
cells present in forewing; marginal cell considerably exceed-
ing third submarginal cell; distal part of wings strongly pa-
pillate, basal part sparsely hairy; jugal lobe of hindwing no
more than one-third as long as vannal lobe; second abscissa
of M + Cu sometimes absent, always shorter than oblique
cu-v and less than one-half as long as M. Mesotibial spur
bifid or multi dentate at apex; tarsal claws with large inner
basal lobe or tooth; tarsal arolia absent (except Cteniosche-
lus).

Female pygidial plate often poorly defined; prepygidial
fimbria absent; female sternum 6 with longitudinal median
carina. Male tergum 7 bilobate or bidentate at apex; gono-
stylus squamiform in dorsal view.

The tribal name Ericrocini was first proposed by Cockerell
and Atkins (1902) to include the Nearctic genus Eric rods.
Ctenioschelini was first introduced by Michener (1965). Even
though Ctenioschelus is an older generic name than Ericrocis,
the Law of Priority applies to tribal names and Ericrocini
thus remains the correct name for this group.

DISCUSSION

The affinities of the Ericrocini have been obscure, though
they have been presumed to be derived from the Centridini
w'hich are their hosts, perhaps via the same stock from which
Epicharis is derived.

CLABISTIC ANALYSIS

The proboscis is considered to be directed downward so that
it has anterior and posterior surfaces, thus Figures 70 and 72
show' the posterior surface of the labiomaxiilary complex.

The cladogram was made with aid of the computer pro-
gram PAUP (Swofford, 1984). Caenonomada, which is un-
doubtedly the centridine genus with the most ancestral traits,
was considered as the outgroup using ordered, unweighted
character (i.e., 0 = primitive, 1 = derived, 2 or more = more
derived characters of a transformation series). Polarity de-
cisions were also decided considering “primitive” antho-
phorids such as Exomaiopsini and in some cases short-tongued
bees. Variables which exhibit two or more characters within
a taxon were scored as the most primitive character found
in that taxon. For example in variable 33 not all species of
Mesonychium have flattened setae, on their meso- or meta-
distitarsus, so the absence of these setae, which is a plesio-
morphy, was used for Mesonychium in the cladistic analysis.
The reasoning is that the plesiomorphic, rather than the apo-
morphic, character will best show the relationships to other
taxa. Therefore the characters of the most primitive members
would be more useful in elucidating cladogenesis.

Table 1 is a list of 67 variables relevant to the Ericrocini,
Rhathymini, and Centridini. Polarities of variables were as-
certained by consideration of the Centridini, from which the
ericrocine bees were presumably derived. Table 2 gives the
raw data. The Centridini are solitary, nest-making bees. Be-
fore 1944 the Centridini had often also included exoma-
lopsine bees, but since Michener (1944), the Centridini has
included only Centris and Epicharis. Snelling (1984) elevated
Ptilotopus, previously a subgenus of Centris, to generic stand-
ing. For purposes of outgroup comparisons we are including
a fourth genus Caenonomada, in the Centridini. Caenono-
mada is the most “primitive” centridine bee, having pre-
viously been placed in the Exomaiopsini (Michener and
Moure, 1957). The reasons for this transfer will be given in
a subsequent paper. The Centridini may be paraphyletic.
Centris, Epicharis, and Ptilotopus share some apomorphies
with the Ericrocini, which are not shared with Caenonomada,
such as fusion of gonostylus with apex of gonocoxite, loss of
arolia, the elongate, narrowed fiabellum which has a cobble-
stone-like posterior surface, profile of the scuteilum vertical
to overhanging the metanotum, profile of the metanotum
more or less vertical, the elongate mesocoxae, the stigma not
extending into the marginal cell and not wider than the pre-
stigma (measured to the costal margin of the wing), and alar
papillae large and not ending in hairs [except Epicharis ( Epi -
charoides) and E. ( Epicharitides ) which have alar papillae
small and ending in hairs]. Therefore Caenonomada may be
the sister group to the rest of the Centridini and Ericrocini.
For purposes of this study we consider Centridini paraphy-
letic.

The Rhathymini are shown as the sister group to Ericrocini
(Fig. 78a) or as the sister group to Ericrocini plus Centridini
(Fig. 78b). Rhathymus shares with Caenonomada and/or
Epicharis plesiomorphic hairy wings, small alar papillae end-
ing in hairs, a hairy propodeal triangle, presence of arolia,
presence of basal bullae on labrum, and unmodified meso-
tibial spurs, all of which are not shared with Ericrocini. The
many common synapomorphies of the Rhathymini and Er-
icrocini could be convergences somewhat reminiscent of con-
vergences between ericrocine and nomadine or melectine

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 3

Table 1. List of variables. Apomorphies are discussed first under each character (plesiomorphies are in brackets; their symbols for Table 2
are 0).

1. Labrum with preapical ridge or tubercle (1). [Labrum simple.]
This ridge is not found in Centridim and is an obvious apo-
morphy of Ericrocini.

2. First flagellar segment of female differentiated, longer than oth-
ers and shorter to longer than scape (1). [First flagellar segment
of female not differentiated, more or less similar in length and
shape to following segments.] A non-differentiated first flagellar
segment is a plesiomorphy for bees in general and is typical of
sphecoid wasps, short-tongued, and exomalopsine bees. Prim-
itive centridine bees possibly had a slightly differentiated first
flagellar segment as seen in Caenonomada. The highly derived
centridines have a very long first flagellar segment. If such Cen-
tridini are ancestral to Ericrocini, their undifferentiated first fla-
gellar segment is apomorphic. If Centridini is monophyletic
(sensu Hennig), i.e., the sister group to Ericrocini, the ericrocine
first flagellar segment would be primitive.

3. Inner eye margins divergent above ( 1 ). [Inner eye margins more
or less parallel.] In nearly all Ericrocini and in all Centridini the
inner eye margins are more or less parallel. The divergent eye
margins apparently have appeared twice, once in Ericrocis and
once in the Mesonychium group.

4. Male flagellar segments greatly elongated, flagellum longer than
body (1). [Male flagellar segments normal, length of a segment
about equal to width.] The bizarre greatly elongated flagellum
of Ctenioschelus, reminiscent of long-homed beetles, is an ob-
vious apomorphy.

5. Mandible of female simple, without subapical tooth (pollex) on
upper margin (1). [Mandible of female with subapical tooth on
upper margin.] Typically bees have toothed mandibles; loss of
this tooth is derived.

6. Paraglossa two-thirds as long as prementum or longer ( 1 ). [Para-
glossa less than two-thirds as long as prementum.] Short para-
glossae are found in short-tongued and exomalopsine bees and
are considered ancestral. Centridini as well as ericrocine bees
have short paraglossae except the Hopliphora group.

7. Stipes without comb ( 1 ). [Stipes with comb.] Most non-parasitic
long-tongued bees have stipital combs. Almost all parasitic an-
thophorids have lost the comb but have retained the preapical
concavity which housed the comb of bristles. The presence of
the comb is a plesiomorphy and its loss an apomorphy asso-
ciated with parasitic habits.

8. Lower end of anterior conjunctival thickening [=suspensory
thickening of Winston (1979)] near basal third of prementum
( 1 ). [Lower end of anterior conjunctival thickening near base of
prementum.] Colletid and andrenid bees have the lower end of
the anterior conjunctival thickening near the basal third of the
prementum. This is the primitive character for Apoidea, where-
as halictids and nearly all long-tongued bees have the lower end
of the anterior conjunctival thickening near the base of the pre-
mentum. Acanthopus has presumably reverted (for Apoidea) to
the ancestral condition which is an apomorphy for Ericrocini.

9. First segment of labial palpus less than twice as long as second
(1). [First segment of labial palpus at least twice as long as
second.] Short-tongued bees have the first and second labial
palpal segments nearly the same length. The evolutionary trend
has been toward a shortening of the second segment and/or
lengthening of the first segment either of which is derived for

Apoidea. In the Ericrocini and Centridini the comparatively
short first segment of the labial palpus is found only in Acan-
thopus and is an apomorphy as is the apomorphy of charac-
ter 8.

10. Mentum appearing Y-shaped because of deep apical emargi-
nation (Fig. 73) (1). Mentum appearing U-shaped because of
deeper emargination (Fig. 71) (2). [Mentum with little if any
apical emargination.] The mentum of most long-tongued bees
is long, narrow and apically with little or no emargination. The
deep apical emargination of the mentum of the Ericrocini is
characteristic for this tribe (Fig. 73) and Nomada (Nomadinae)
although in Mesopiia the emargination is not strong (Fig. 76).
Acanthopus has the most derived mentum in that it is so deeply
divided that it appears U-shaped.

1 1 . Lorum basally divided such that the loral apron is separate
sclerites held together by membrane (1). [Lorum V-shaped.] The
lorum in Centridini as well as other anthophorids is V- or Y-
shaped (Fig. 73). Acanthopus has a uniquely apomorphic lorum,
basally divided such that the loral apron (Michener, 1985) is
two sclerites held mediobasally by membrane and is fused to
the cardines (Fig. 71).

12. Postflabellum present (1). [Postfiabellum absent.] The postfia-
bellum (Michener and Brooks, 1984) is unique to the ericrocine
bees and is an obvious apomorphy.

13. Forewing tips (and sometimes marginal cell) infuscated with
rest of wing clear (1). [Forewing tips concolorous with rest of
wing, clear or infuscated.] Infuscated wing tips have appeared
convergently many times in the Apoidea. The primitive con-
dition of a concolorous wing, whether infuscated or clear, is
found in most bees including the Centridini and most Ericrocini.
Only the Ctenioschelus group and some Mesopiia have infus-
cated wing tips although they are variable in Ctenioschelus since
the Middle American populations have less distinctive infus-
cation than the Brazilian ones.

14. Maxillary palpus with three or four segments (1). Maxillary
palpus with one or two segments (2). Maxillary palpus absent
or represented by a small bump fused to stipes (3). [Maxillary
palpus with five or six segments.] The maxillary palpus primi-
tively has six segments as seen in most bees including Caenon-
omada. Other centridines have five segments. The evolutionary
direction in the ericrocines has been reduction and sometimes
fusion of segments, culminating in complete loss of the palpus
in Acanthopus.

15. Metasomal integument with metallic reflection (1). [Metasomal
integument without metallic reflection.] Note that this statement
refers to the color of the integument, not the metallic body hairs.
The plesiomorphic condition is found in most Centridini [Cen-
tris s. str., some C. ( Paracentris ), and C. ( Wagenknechtia ) have
metallic terga] and almost all Ericrocini where the integument
is black to red-brown with no metallic reflections. Only Abro-
melissa has the apomorphic metasomal integumental color which
is metallic blue.

16. Vestiture metallic in color (1). [Vestiture non-metallic.] The
primitive condition is found in Centridini and Ericrocis. All
other ericrocines have metallic vestiture.

1 7. Profile of scutellum vertical or at least at strong angle to scutum
( 1 ). Profile of scutellum overhanging metanotum (2). [Profile of

4 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

Table 1. Continued.

scutellum with posterior part more or less horizontal to a 45°
angle.] Most of the short-tongued and exomalopsine bees have
the primitive type of scutellum as does Caenonomada. The
ericrocir.es. like the rest of the centridines, have an apomorphic
scutellum whose profile is vertical and additionally overhangs
the metanotum in Mesoplia, Abromelissa, and the Ctenioschelus
group.

18. Profile of metanotum vertical (1). [Profile of metanotum slant-
ing, more or less at a 45° angle.] Most short-tongued bees, all
exomalopsine bees and Caenonomada have the plesiomorphic
type of metanotum which slants. All the rest of the centridines
and ericrocin.es have an apomorphic, more or less vertical meta-
notum.

19. Propodeal triangle hairless (1). [Propodeal triangle hairy.] A
propodeal triangle with hair is found in all Centridini. The Er-
icrocini have a hairless propodeal triangle which is presumably
an apomorpfay, although many other bee groups possess this
character.

20. Propodeum in profile with horizontal basal zone short, less than
one-third as long as declivous surface (1). Propodeum in profile
entirely declivous (2). [Propodeum in profile with horizontal
basal zone long, about two-thirds as long as declivous surface.]
Most “primitive” anthophorids and Caenonomada have a pro-
podeum with a long horizontal basal zone. The evolutionary
trend has been a shortening of this zone.

21. Jugal lobe shortened, one-third to one-half as long as vannal
lobe measured from wing base (1). Jugal lobe short, one-fourth
to less than one-third as long as vannal lobe (2). Jugal lobe very
short, less than one-fourth as long as vannal lobe (3). [Jugal lobe
long, apex much nearer vannal incision than wing base.] The
evolutionary direction has been a shortening of the jugal lobe
from a plesiomorphic long lobe of about three-fourths the length
of the vannal lobe (as in Caenonomada) to a smaller one.

22. Hindwing with second abscissa of M+Cu almost absent to half
the length of the crossvein cu-v (1). [Hindwing with second
abscissa of M+Cu about equal in length to crossvein cu-v, sec-
ond abscissa of M + Cu 0.75 to less than 1.50 times as long as
crossvein cu-v.] In the Centridini the second abscissa of the M+Cu
is about equal to the length of vein cu-v. The evolutionary trend
in the Ericrocini has been toward a shortening of the second
abscissa of the M+Cu which has culminated in its near to com-
plete absence.

23. Hindwing with second abscissa of M+Cu considerably shorter
than M (1). [Hindwing with second abscissa of M + Cu about as
long as vein M.] This character is correlated with the previous
character because as the second abscissa of M + Cu shortens
(which is primitively almost equal in length to vein M as seen
in short-tongued and exomalopsine bees), vein M will obviously
be lengthened.

24. Hindwing with crossvein cu-v slanted toward wing base from
second abscissa of M+Cu (1). [Hindwing with crossvein cu-v
perpendicular to slanted apically to wing tip from second ab-
scissa of M + Cu.] The hindwing of short-tongued bees generally
has vein cu-v perpendicular to or slanted apically to wing tip
from the second abscissa of vein M + Cu. Generally in the de-
rived anthophorids this crossvein has become slanted toward
wing base from the second abscissa of M + Cu.

25. Stigma not or scarcely extending into marginal cell, not wider
than prestigma measured to costal margin of wing (1). [Stigma

slender, posterior margin angulate at base of vein R, extending
into marginal cell but oblique and straight there, not or little
wider than prestigma.] Caenonomada has a moderately large
stigma that extends into the marginal cell. A slendering of the
stigma and its decreasing extension into the marginal cell is the
derived condition. Caenonomada has a stigma intermediate be-
tween most exomalopsines and the rest of the centridines.

26. Stigma about as long as prestigma (1). Stigma shorter than pre-
stigma, absent to nearly absent (2). [Stigma longer than prestig-
ma.] The plesiomorphic long stigma as seen in Caenonomada
has apomorphically decreased in length until it is nearly absent
in the other Centridini, but it has decreased in length somewhat
less in the Ericrocini.

27. Wings bare or with small patches of hair (1). [Wings hairy
throughout or at least over large areas.] Hairy wings are found
in the Exomalopsini, Melitomini, Eucerini, Caenonomada, and
Epicharis. The Ericrocini have lost the hair.

28. Alar papillae large, not ending in hairs (1). [Alar papillae small
and ending in hairs.] “Primitive” anthophorids, Caenonomada,
and two subgenera of Epicharis ( Epicharoides and Epicharitides)
have alar papillae that end in hairs. In the other Centridini and
the Ericrocini the papillae have become larger and have lost the
small apical hairs.

29. First recurrent vein distad to interstitial, or nearly so with 1st
transverse cubital vein (1). [First recurrent vein intersecting dis-
tal half of posterior margin of submarginal cell 2.] The first
recurrent vein intersects the distal half of the posterior margin
of submarginal cell 2 in all Centridini. The interstitial first re-
current and 1st transverse cubital veins in Ericrocini is an apo-
morphy. The polarity of this character is reversed for the Apoi-
dea since exomalopsine, melitomine, and eucenne bees have
the first recurrent and 1st transverse cubital interstitial.

30. First recurrent vein (as well as second) intersecting submarginal
cell 3 (1). [First recurrent vein basad to interstitial with 1st
transverse cubital vein.] In almost all apoids the first recurrent
vein intersects submarginal cell 2 or is interstitial with the vein
shared by submarginal cells 2 and 3. It is extremely rare to find
the first and second recurrent veins entering submarginal cell 3.
This is an obvious apomorphy and unique to Acanthopus among
the ericrocine-centridine bees.

31. Marginal cell short, length less than 1.2 the distance from apex
of marginal cell to wing tip (1). Marginal cell very short, length
less than 0.7 the distance from apex of marginal cell to wing tip
or marginal cell very long, length 1.8 the distance from apex of
marginal cell to wing tip (2). [Marginal cell long, length 1 .2-1 .45
the distance from apex of marginal cell to wing tip.] Primitively
the marginal is long, longer than the distance from the apex of
the marginal cell to the wing tip. This plesiomorphy is seen in
exomalopsine bees as well as Epicharis and Caenonomada. This
Y-shaped variable consists of an apomorphic shortening in the
Ericrocini starting from a long marginal cell, but in Acanthopus
the evolutionary direction apparently changed to a lengthening
of the marginal cell.

32. Mesobasitarsus laterally compressed, posterior margin carinate
(1). [Mesobasitarsus somewhat round in cross section to slightly
compressed, posterior margin not carinate.] The posterior mar-
gin of the mesobasitarsus is commonly compressed and carinate,
terminating in a blunt tooth or spur that extends beyond the
end of the segment. Its presence, found in most ericrocines, is
an apomorphy since the Centridini lack it.

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 5

Table 1. Continued.

33. Meso- and/or metadistitarsi with flattened setae ( 1 ). [Meso- and/
or metadistitarsi normal, without flattened setae.] The dark flat-
tened setae of the meso- and/or metadistitarsi have presumably
arisen twice in the Ericrocini. Ericrocis and most Mesonychium
have this apomorphy not found in the Centridini.

34. Metadistitarsus five times longer than wide ( 1 ). [Metadistitarsus
at most four times longer than wide.] The greatly elongated
metadistitarsus of Acanthopus is unique in the Ericrocini and
Centndini.

35. Arolia absent (1). [Arolia present, small.] The primitive centri-
dine Caenonomada has small arolia. Arolia have been lost in
all other centridines and ericrocines except in Aglaomelissa and
Ctenioschelus in which they are small. The loss of arolia is
certainly an apomorphy since almost all bees and sphecoid wasps
have arolia. If the cladogram is correct it is very unlikely that
the presence of arolia in Aglaomelissa and Ctenioschelus is in-
dependently derived since the arolium is a complex structure,
composed of many parts, all of which are present in these two
as in other aroliate bees. Moreover, it would be improbable if
their presence requires five independent losses of arolia in the
other Ericrocini. We believe that, assuming the accuracy of the
cladogram, the origin of arolia in Aglaomelissa and Cteniosche-
lus is best explained as a reversion due to reactivation of genes
that were suppressed during much of centridine and ericrocine
evolution.

36. Mesotibial spur modified with several apical teeth (1). [Meso-
tibial spur normal, apically simple.] The modified mesotibial
spur of the Ericrocini is an obvious apomorphy which the Cen-
tridini do not possess.

37. Females with tooth of tarsal claw a flattened basal lobe (1).
[Female with tooth of tarsal claw not a flattened basal lobe.]
The modified tooth becoming a flattened lobe is an apomorphy
of Ericrocini which has convergently appeared at least three
times in the Anthophoridae. Almost all non-parasitic bees have
the plesiomorphic toothed claw, whereas all Ericrocini, Rhath-
ymini, Melectini, and almost all Nomadinae have the tooth a
flattened basal lobe.

38. Strigilar malus without teeth (1). [Strigilar malus with teeth.]
Primitively the strigilar malus has teeth as found in many short-
tongued bees. The loss of the teeth has occurred twice in the
Ericrocini, once in the Mesonychium group and again in Agla-
omelissa. This character may be of little phylogenetic signifi-
cance since it is so variable throughout the Anthophoridae but
it further unites the Mesonychium group and may prove valuable
for future studies.

39. Scopa absent (1). [Scopa present.] All non-parasitic female bees
have scopae [except Hylaeinae (Colletidae)]. The loss of the
scopa is an obvious apomorphy seen in the Ericrocini and to
varying degrees in all parasitic bees.

40. Anterior mesepistemal carina present (1). [Anterior mesepi-
stemal carina absent.] The junction of the short anterior face and
the long lateral face of the mesepistemum is rounded in Cen-
tridini. The apomorphic state has apparently appeared twice,
once in Acanthopus and again in the Ctenioschelus group though
lacking in Ctenioschelus.

4 1 . Stemopleural ridge present ( 1 ). [Stemopleural ridge absent.] The
presence of the stemopleural ridge is an apomorphy found only
in the Hopliphora group.

42. Supraspiracular ridge well developed, terminating abruptly in a
blunt tooth-like process (1). [Supraspiracular ridge weak to ab-
sent.] The supraspiracular ridge is not developed in Centridini
and many ericrocines. It is developed in the Mesonychium and
Ctenioschelus groups (except Aglaomelissa).

43. Scutellum bilobed (1). Scutellum bilobed and projecting over
propodeum (2). [Scutellum not projecting over propodeum, sim-
ple.] In Centridini as in most non-parasitic anthophorids the
scutellum is not modified. The scutellum is commonly modified
in most parasitic anthophorids and all Ericrocini. The modifi-
cation in ericrocine bees is the presence of two rounded lobes
which are further modified in the Ctenioschelus group (except
Aglaomelissa) as dorsoventrally flattened plate-like eminences
extending over the vertical metanotum and propodeum.

44. Midcoxal length 1.00-1.35 length of distance from summit of
coxa to hind wing base (1). Midcoxal length at least 1.36 length
of distance from summit of coxa to hind wing base (2). [Midcoxal
length 0.85-0.99 length of distance from summit of coxa to hind
wing base.] In many parasitic anthophorine bees the midcoxa
has become elongated as well as more exposed and hence is a
further derivation as seen in most ericrocines (except the re-
version found in the Mesonychium group) and centridines (ex-
cept Caenonomada).

45. Hind basitarsus of female with distal process, not provided with
a pencillus (1). [Hind basitarsus of female with distal process
ending in a pencillus.] Almost all non-parasitic anthophorid bees
have a pencillus. The loss of the pencillus in almost all parasitic
bees is an apomorphy.

46. Basitibial plate absent or only represented by a carina (1). [Ba-
sitibial plate of female present, well developed.] The basitibial
plate present in female Centridini has been lost in the Ericrocini
as well as most other parasitic bees and is an apomorphy.

47. Basitibial plate of male incomplete, represented by a scale or
series of tubercles (1). Basitibial plate absent or only represented
by a carina (2). [Basitibial plate of male present, well developed.]
Male Centridini have well-developed basitibial plates (except
Centris). The loss of these plates in male Ericrocini is an apo-
morphy.

48. Dorsum of metasomal tergum 1 subangulate in profile (Fig. 77)
(1). [Dorsum of metasomal tergum 1 rounded in profile (Fig.
76).] Usually, the vertical and horizontal faces of the first meta-
somal tergum, as viewed in profile, meet in a distinctly rounded
angle, the junction, however, may be so narrowly and abruptly
rounded as to appear angulate. This is an obvious apomorphy
of the Ctenioschelus group.

49. Prepygidial and pygidial fimbriae absent (1). [Prepygidial and
pygidial fimbriae present.] The Centridini as well as all non-
parasitic anthophorine bees have prepygidial and pygidial fim-
briae. The loss of these fimbriae in the Ericrocini is derived.

50. Metasomal terga with patches of appressed white hair ( 1 ). [Meta-
somal terga, without patches of appressed white hair.] The Cen-
tridini have the metasoma with long, erect to suberect hair and/
or entirely covered with appressed hair or virtually all bare.
Most ericrocines have patches of white hair either laterally on
the terga or in various patterns, but this apparently has been
reversed in the Hopliphora group and Abromelissa. This apo-
morphy has arisen many times in other parasitic anthophorids.

6 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

Table 1. Continued.

5 1 . Metasomal sternum 3 of male with dense lateral patch of white
hair (1). [Metasomal sternum 3 of male simple, without dense
lateral patch of white hair.] The Centridini lack the dense lateral
patch of white hair on metasomal sternum 3. All Ericrocini have
this apomorphy except the Hopliphora and Mesonychium groups.

52. Metasomal sterna 4 and/or 5 of male with dense apical band of
long curved hair ( 1 ). [Metasomal sterna 4 and 5 of male simple
without apical band of long curved hair.] Caenonomada. Epi-
charis, and the Ericrocini (except Mesoplia and Mesonychium)
share the apomorphy of an apical band of long curved hair on
metasomal sterna 4 and/or 5. “Primitive” anthophorids such
as Exomalopsini, Melitomini, and Eucerini lack bands.

53. Metasomal sternum 5 (and sometimes 4) on basal half of disc
with felt-like pad of dense hair (1). [Metasomal sternum 5 and
4 simple, without felt-like pad of dense hair.] The presence of
a felt-like pad of short pubescence on metasomal sternum 5 (and
sometimes 4) which Centridini lacks is an obvious apomorphy
found in all Ericrocini except the Mesonychium group.

54. Male metasomal sternum 4 concealed (1). [Male metasomal
sternum 4 normal, not concealed.] Because the margins of the
fourth metasomal sternum may be broadly incurved, each seg-
ment may be largely hidden under the preceding segment. This
apomorphy is found scattered throughout the Ericrocini.

55. Male metasomal sternum 5 concealed (1). [Male metasomal
sternum 5 normal, not concealed.] See discussion for character
54.

56. Female metasomal sterna 2-5 longitudinally carinate medially
(1). [Female metasomal sterna 2-5, not carinate medially.] Fe-
male ericrocines usually have metasomal sterna 2-5 uniformly
flat as in most bees. Mesocheira is the only genus which has the
unique apomorphy of sterna 2-5 carinate medially.

57. Female metasomal sternum 6 with longitudinal median carina
( 1 ). [Female metasomal sternum 6 longitudinally simple without
median carina.] All Ericrocini have this unique apomorphy which
the Centridini as well as other non-parasitic anthophorids lack.

58. Sixth metasomal sternum of male medioapically flat, not con-
vex, usually entire or emarginate, rarely pointed ( 1 ). [Sixth meta-
somal sternum of male bluntly pointed medioapically, strongly
convex.] The sixth metasomal sternum of male medioapically
pointed and convex is seen in all exomalopsine, melitomine,
eucerine, and centridine bees.

59. Sternal apodemes of some metasomal sterna without dorsal pro-
cesses (1). [Sternal apodemes of metasomal sterna with dorsal
processes.] Centridini have metasomal sternal apodemes with
dorsal processes. This condition is commonly found among the
non-parasitic Anthophoridae. All Ericrocini have lost the dorsal
processes except Ericrocis. This loss is an apomorphy.

60. Eighth metasomal sternum of male without or with greatly re-
duced spiculum (1). [Eighth metasomal sternum of male with
spiculum.] Most anthophorid bees have a spiculum as found in
the Centridini. An obvious apomorphy is loss or great reduction
of the spiculum as seen in the Ericrocini except Abromelissa.

61. Spatha short, weakly developed (1). Spatha absent (2). [Spatha
present, well developed.] Centridini except Epicharis have a well
to weakly developed spatha. The presence of a spatha is pre-
sumably a plesiomorphy since it is commonly found among
“primitive” anthophorids. Apparently the loss of the spatha in
Ericrocini is an apomorphy.

62. Eighth metasomal sternum of male without apical process (1).
[Eighth metasomal sternum of male with apical process.] Many
short-tongued bees, the "primitive” anthophorid and Centridini
have a well-developed apical process which may be variously
modified. The reduction of this apical process such that the disc
of the eighth sternum is broader than long is an apomorphy
typical of Ericrocini.

63. Eighth metasomal sternum of male a broad plate (1). [Eighth
metasomal sternum of male short, transverse, sometimes with
one or two long apical processes.] For discussion see variable
62.

64. Penis valve without basolateral lobe (1). [Penis valve with ba-
solateral lobe.] The penis valves of centridine bees have basolat-
eral lobes. The presence of these lobes is an apomorphy though
a weak one since it has apparently appeared three times, being
a character of Ericrocis, Aglaomelissa, and the Hopliphora group.

65. Gonostylus of male greatly reduced to absent, if present then
represented only by a membranous, flat, circular to slit-like area
(1). [Gonostylus of male an elongate appendage.] The presence
of a gonostylus as an elongate appendage often incurved apically
distad of the penis valves is common in the “primitive” an-
thophorids and the Centridini. Recognition of the gonostylus is
often aided by the long setae which all anthophorid bees have
on their gonostyli, as well as the reduced sclerotization of the
gonostylus. The most primitive condition is articulation to the
apex of the gonocoxite as it is in Caenonomada. In the other
centridines and ericrocines it is fused to the apex of the gono-
coxite and is very narrow, round in cross section and more
elongate. The reduction or absence of the gonostylus is a weak
apomorphy since it has disappeared in three separate ericrocine
groups.

66. Mesopleuron with knob-like process (1). [Mesopleuron without
knob-like process.] The knob-like process on the mesopleuron
of Rhathymus is an obvious autoapomorphy.

67. Apical margin of metasomal sternum 5 laterally with notch (1)
(Fig. 76). [Apical margin of metasomal sternum 5 laterally sim-
ple.] This unique autoapomorphy is found only in Mesocheira.

bees. Figure 78b necessitates 1 1 convergences between the
two tribes, which is possible considering there are 12 con-
vergences [characters 7, 17, 18, 20(2), 21(2), 37, 39, 45-47,
49, 58, and 59] between the melectine and the ericrocine-
rhathymine bees. Nevertheless, the male terminalia of Eri-
crocini and Rhathymini are similar and in conjunction with

the numerous apparent synapomorphies shared by the two
tribes, we prefer Figure 78a. The cladogram of Figure 78a
eliminates all 1 1 convergences of Figure 78b and so is 11
steps shorter.

Unique characters for taxa were obvious apomorphies,
whereas other polarity decisions encompassed centridine

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 7

Table 2. Matrix of characters listed in Table 1. Variables 1-67 are represented by the columns from left to right. Variables which are not
constant are indicated by both characters which appear in the genus, i.e., 0.

CAENONOM

RHATHYMU

MESOPLIA

HOPLIPHO

MESONYCH

ABROMELI

ERICROCI

AGLAOMEL

CTENIOSC

MESOCHEI

ACANTHOP

0000000000000000000 12011 000000000000000000000000000 1 00000000 1 000000
010010101 0000300 110230111 00020000000 1 0 1 000000 1201001 000000 1 0000 1010
1 100001001010101211221111111201100111010001011201110101011112110000
1100011001010101111221111111201100111010101111201001100011112111000
1 1 10001001010201111221111111201 100111110011011201100010011112110100
1110000001010111211231111111201100111110011011201001000011102110000
1110001001010100111221111111202110111010001211201111100011012111100
0100101001011201211231111111201000011111001111211111100011112111000
1101101001011201211231111111201000011010012111211111100011112110000
0100101001011201211231111111201100111011012111211111101111112110001
1100111112110301111221111111210001111011101111201001101011112111000

outgroup comparisons or centridine-exomalopsine outgroup
comparisons or comparisons against short-tongued bees, with
the variables of the ericrocine bees. The long first flagellar
segment of the Centridini, which is a derived feature for the
family Anthophoridae, is considered here as a plesiomorphy,
the somewhat non-differentiated first flagellar segment of the
ericrocines being the apomorphous character.

Table 1 includes notes on the distribution of characters of
the variables and the bases for our judgments of polarity
(evolutional direction). Discussion is frequently abbrevi-
ated but is sufficient to suggest reasons for our decisions.

DISCUSSION

Linsley ( 1939) suggested that Ericrocis may have arisen from
Centris ( =Hemisia ) or Epicharis. Michener (1944) stated that
Melectini, Ericrocini, and the Rhathymini may have had a
common origin with Anthophora but later he (Michener,
1974) separated the melectine and rhathymine-ericrocine lin-
eages in his dendrogram, indicating common origins of the
Anthophorini with the former and Centridini with the latter.
Our work supports Michener’s latter hypothesis and is more
or less in agreement with Rozen’s larval work (1969). Rozen
states that the larvae of melectines, ericrocines, and rha-
thymines share an apomorphy not found in centridine or
anthophorine larvae, i.e., they lack galeae. But as he points
out, this may not indicate a common origin for these parasitic
bees since galeae have been lost in many unrelated groups
of bees and loss characters are often convergent. Rozen fur-
ther states that the Melectini share a plesiomorphy with the
Anthophorini not shared with the Ericrocini and Rhathy-
mini; i.e., the maxillary palpus is preapical in position. The
Ericrocini and Rhathymini have maxillary palpi which are
apical as well as greatly elongated labiomaxillary regions.
These apomorphies support the contention that the Ericro-
cini and Rhathymini are sister groups. On the other hand
the strongly denticulate atrial wall and spinous primary tra-
cheal opening of the ericrocines (not found in rhathymines)
is similar to the spiracular structure of Epicharis and an-
thophorines and therefore could suggest a separate origin for
the Rhathymini. We believe that the rhathymine-ericrocine

clade is monophyletic ( sensu Hennig) since it has 10 adult
and two larval synapomorphies. It is unlikely that all of the
apomorphies are convergent and have appeared de novo in
each tribe. We have presented, however, other characters
which do not support this hypothesis.

We have recognized several lineages within the Ericrocini:
(a) Ericrocis group ( Ericrocis ), (b) Hopliphora group ( Hopli -
phora and Acanthopus), (c) Mesoplia group (Mesoplia), (d)
Mesonychium group ( Mesonychium and Abromelissa), and
(e) Ctenioschelus group ( Ctenioschelus , Aglaomelissa, and
Mesocheira). Here and below the numbers in parentheses
represent variables as numbered in Table 1 . The most “prim-
itive” lineage is presumably Ericrocis since it lacks metallic
setae (16) and integumental coloration (15) and has meta-
somal sternal apodemes (59) like Centridini. This is not fully
convincing since the loss of metallic coloration could be a
reversion in Ericrocis rather than a plesiomorphy shared with
Centridini. Metallic hair has arisen three times at least in the
Anthophorinae, in Amegilla (Anthophorini), in Thyreus
(Melectini) as well as in the Ericrocini. Many Thyreus have
non-metallic white appressed vestiture like Ericrocis and a
polarity decision of whether that type of vestiture is primitive
or derived will probably be similarly uncertain.

Acanthopus has the most derived mouthparts and legs of
any ericrocine and two unique apomorphies in the forewing,
making its relationship to the rest of the Ericrocini obscure.
Nevertheless, it shares three strong apomorphies with Hop-
liphora, i.e., relatively long paraglossae (6), presence of the
stemopleural ridge (4 1 ) and the absence of tergal patches of
white appressed hair (50). Hopliphora and Acanthopus are
the largest ericrocines and both have large centridine hosts.
Lack of apomorphies for Hopliphora suggests that it is pos-
sibly a paraphyletic group from which Acanthopus evolved.
We have no problem in recognizing paraphyletic taxa since
we find it unnecessary that classifications and cladograms be
redundant, especially when more than one cladogram is
equally plausible (or parsimonious).

The Mesoplia, Mesonychium, and Ctenioschelus groups
have three apomorphies, a well-developed scutellum which
overhangs the metanotum (43) (reverted to a less developed
state in Mesonychium ), metasomal sternum 3 of male with

8 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

dense lateral patch of white hair (51) (which is reversed in
the Mesonychium group), and penis valve with basolateral
lobe (64) (lost in Aglaomelissa). In view of the reversals, one
must admit that the three characters are not very convincing.

Mesoplia is quite distinctive, having elongate segments in
the maxillary palpus (14), the male has lost the apical bands
of long curved hair on metasomal sterna 4 and 5 (52) and
has a broadly emarginate sternum 5 such that it is concealed
under sternum 4 (55).

The Mesonychium and Ctenioschelus groups share a re-
duction of the number of maxillary palpal segments to one
or two (14) (reduced to three in Abromelissa), a reduction of
the length of the jugal lobe to less than one-fourth the distance
between the vannal incision and wing base (21) (reduced to
between one-third and one-fourth this distance in Mesony-
chium), and presence of the supraspiracular ridge (42).

The Mesonychium group is characterized by five strong
apomorphies. The inner eye margins are divergent above (3),
the strigiiar malus lacks teeth (38), the midcoxa is short (44),
metasomal sternum 3 lacks a dense lateral patch of white
hair (51), and sterna 4 and 5 lack dense felt-like pads of
pubescence (53) which ancestrally cover the basal halves of
their discs. It is interesting also that Abromelissa is the only
ericrocine genus with metallic integument (15). The Meso-
nychium group, although it includes Amazonian species, is
largely peripheral to the distribution of other ericrocine gen-
era. Mesonychium is one of the largest genera in the tribe,
with about half a dozen species in Chile and temperate Ar-
gentina. Also limited to Chile is its possible derivative, Abro-
melissa, so Mesonychium may be paraphyletic.

The Ctenioschelus group is distinguished by several de-
rived characters: the subapical mandibular tooth is lost (5),
the forewing tips are infuscated (13), and tergum 1 has a
relatively sharply angulate profile (48) (Fig. 76). Other note-
worthy characters which are found in two of the three genera
are the presence of arolia (35) (except Mesocheira), the api-
cally simple labrum (1), the presence of an anterior mes-
epistemal carina (40) (except Ctenioschelus), and the projec-
tion of the scutellar lobes over the propodeum (43) (except
Aglaomelissa). There is also a trend toward the development
of the occipital margin into a sharp ridge or flange, or a flange-
like carina across the entire pronotum, and of strongly de-
veloped acetabular carinae with the procoxae deeply re-
cessed. The Ctenioschelus group is centered in Amazonia but
Mesocheira ranges from central Mexico to Paraguay. Cte-
nioschelus and Aglaomelissa are primarily South American,
but both range north to Costa Rica.

KEY TO GENERA OF ERICROCINI

la. Second and third submargina! cells of forewing each re-
ceiving a recurrent vein; metatarsus without dense fringe

of long, dark plumose hairs 2

b. Third submarginal cell large, receiving both recurrent
veins; metatarsus very long, with dense brush of long,

dark, plumose hairs Acanthopus

2a. Juncture of basal and discal surfaces of tergum 1 slightly
humped and subangulate in middle (Fig. 77); mesoba-

sitarsus without distal, flattened, spine-like projection
on posterior margin (Fig. 49); mandible simple .... 3
b. Juncture of basal and discal surfaces of tergum 1 evenly
rounded, never appearing subangulate (Fig. 76); meso-
basitarsus almost always with distal, flattened process
on posterior margin, often continued basad as a raised,
cariniform ridge (Fig. 1); mandible usually with single

preapical tooth 5

3a. Scutellum bituberculate, the processes stout, subconical
and suberect; forewing dusky with apex darker .... 4
b. Scutellum bituberculate, the processes flat and plate-like,
directed caudad; forewing dusky, with an apical cloud

in marginal cell in addition to that at wing apex

Mesocheira

4a. Pronotum carinate between collar and lobe; mesepister-
num with lamelliform ridge between anterior and lateral
surfaces; male antenna normal, not extending much be-
yond tegula Aglaomelissa

b. Pronotum not carinate between collar and lobe, end of
collar clearly defined; mesepistemum abruptly rounded
between anterior and lateral surfaces; male flagellar seg-
ments greatly elongate, flagellum extending well beyond

apex of abdomen Ctenioschelus

5a. Meso- and metadistitarsi with a cluster of appressed,
short flattened setae on each side (Fig. 28) (setae reduced
in some Mesonychium in which antennocular distance
is greater than antennal socket diameter and third sub-
marginal cell is distinctly narrowed anteriorly); inner eye
margins divergent above; male gonostylus without dor-
sal lobe (Figs. 33, 38) 6

b. Meso- and metadistitarsi with normal setae and hairs,
without flattened setae (Fig. 2); inner eye margins var-
ious, but usually not divergent above; male gonostylus
with dorsal lobe (Figs. 7, 22, 43) (except some Hopli-

phora) 7

6a. Abdominal scales contrasting black and white (may be
somewhat tawny); labrum with erect preapical median
tubercle; scutellum without raised, mammiform tuber-
cles Ericrocis

b. Abdominal scales or hairs iridescent blue or green, with
or without contrasting whitish hairs; labrum with trans-
verse preapical ridge; scutellum usually with a pair of

mammiform tubercles Mesonychium

7a. Abdominal integument blackish, usually more or less
covered with blue or green-reflective, appressed, scale-
like hairs, but if not, mesepistemal hairs are black; tegula
rectangular or elliptical but not abruptly narrowed an-
teriorly (Figs. 61, 63, 64) 8

b. Abdominal integument shiny metallic blue, with sparse
erect blackish hairs; hairs of thorax whitish; tegula

abruptly narrower in anterior one-third (Fig. 66)

Abromelissa

8a. Tegula oval, outer margin evenly curved (Fig. 61); ab-
domen usually appearing black, without patterns of ap-
pressed pale pubescence Hopliphora

b. Tegula, from above, approximately elliptical, often with
outer margin somewhat sinuate in part and with pos-
terior margin oblique or truncate; abdomen bright iri-

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Snelling and Brooks: The Tribe Ericrocini 9

descent bluish or greenish, often with conspicuous patches
of white or yellowish pubescence Mesoplia

Ericrocis Group

Although the body is richly marked with conspicuous pat-
terns of blackish and white to tawny pubescence, the com-
plete lack of metallic-reflective hairs or scales is diagnostic
for this group. Other characteristics of the Ericrocis group
are: inner eye margins divergent above (shared with Meso-
nychium group); meso- and metadistitarsi with flattened se-
tae (shared with some Mesonychium ); sternal apodemes of
some metasomal sterna without dorsal processes (unique
within the Ericrocini).

(A) Inner eye margins divergent above. (B) Ocellocular
distance greater than diameter of anterior ocellus. (C) Man-
dible with preapical tooth. (D) Maxillary palpus two-seg-
mented. (E) Hypostomal carina low, uniform. (F) Malus of
protibia short, less than half as long as velum, finely serrate
on inner margin. (G) Mesobasitarsus usually compressed and
more or less sharply carinate along posterior margin; distal
process usually present. (H) Meso- and metadistitarsi with
lateral patch of short, flattened setae on each side (Fig. 28).
(I) Metatarsus without posterior fringe. (J) Marginal cell of
forewing about 0.80 times distance from its apex to wing tip.

(K) Juncture of basal and discal faces of first tergum rounded.

(L) Male seventh tergum bilobate at apex or with two stout
teeth. (M) Gonostylus without dorsal lobe.

The one genus included in this group, Ericrocis , is restrict-
ed to the Nearctic Region.

Genus Ericrocis Cresson

Figures 34-38, 60, 72-74

Ericrocis Cresson, 1887:131, 134. Type species: ? Crocisa
lata Cresson, 1878 (monobasic).

DIAGNOSIS

Abdominal color black, with conspicuous pattern of ap-
pressed, plumose white to somewhat tawny hairs; labrum
with a median preapical tubercle; dorsal face of scutellum
without tubercles.

DESCRIPTION

(1) Head much broader than long; inner eye margins essen-
tially straight, strongly divergent above; occipital margin
nearly flat, slightly elevated above ocelli. (2) Maxillary palpus
two-segmented, first segment distinctly longer than second.
(3) Labrum with median preapical tubercle; apical margin

subtruncate. (4) Interantennal distance greater than antennal
socket diameter; antennocular distance greater than antennal
socket diameter. (5) Ocelloccipital distance slightly greater
than diameter of anterior ocellus. (6) Occipital margin sub-
angulate. (7) Antenna short in both sexes; minimum length
of first flagellar segment about equal to maximum width and
shorter than second segment on same side.

(8) Pronotum not carinate between collar and posterior
lobe. (9) Midline of mesoscutum deeply impressed, more
weakly so distad; parapsidal lines in broad, shallow impres-
sions. ( 1 0) Dorsal face of scutellum broadly impressed along
midline, broadly convex on either side. (11) Anterior mes-
epistemal carina, acetabular carina and stemopleural ridge
absent. (12) Supraspiracular ridge evanescent distad.

( 1 3) Tegula (Fig. 65) rectangular, narrowed anteriorly, out-
er margin somewhat sinuate. (14) Second submarginal cell
narrower on M than first or third; 1 st m-cu interstitial with
1st r-m; 2nd m-cu a little basad of 2nd r-m.

(15) Mesotibial spur slender, parallel-sided, apex bispi-
nose, outer spine often obsolete, intercalary denticles present.

( 1 6) Metatrochanter rounded ventrally; metatibial spurs nor-
mal; metadistitarsus about 2.5 times longer than wide.

(17) Female sixth tergum with weakly defined pygidial
plate, apex narrowly rounded. (18) Male fourth sternum
broadly concave along apical margin, with a distal fringe of
long, dark plumose hairs, their apices broadly reflexed. (19)
Male fifth sternum broadly and more shallowly concave along
apical margin, with or without distal fringe. (20) Male sev-
enth sternum (Fig. 34) transverse, evenly rounded or with
short, truncate projection. (21) Male eighth sternum (Fig. 35)
short, bilobate at apex. (22) Male gonostylus short, thick,
truncate; dorsal lobe absent; inner apical sclerotization of
gonocoxite poorly defined. (23) Penis valve abruptly deflect-
ed ventrad; basolateral lobe absent (Fig. 38).

DISCUSSION

This Nearctic genus is, in many respects similar to the South
American genus Mesonychium, with which it shares the pres-
ence of a cluster of short, flattened setae on either side of the
meso- and metadistitarsi, a feature unique to these two gen-
era. Unlike Mesonychium and all other ericrocine genera,
Ericrocis lacks metallic hairs or scales. Instead, there are
richly marked patterns of black and white and (sometimes)
tawny pubescence. Ericrocis includes two species: E. lata
(Cresson) and E. pintada Snelling and Zavortink. The genus
was revised by Snelling and Zavortink (1985).

It should be noted that the figures of the labiomaxillary
complex cited by Winston (1979: fig. 44) as those of Meso-
cheira bicolor appear, instead, to be based on Ericrocis lata.
Michener and Fraser (1978) figure mandibles purported to

Figures 1-7. Mesoplia (M.) azurea. 1, female mesobasitarsus, pilosity omitted; 2, metadistitarsus, lateral view; 3, male sternum 7; 4, male
sternum 8; 5-7, male genital capsule, lateral, ventral, and dorsal views. Scale line = 1.00 mm (Figs. 3-7 only).

10 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 1 1

Figures 8-12. Mesoplia ( Eumelissa ) decorata, male. 8, sternum 7; 9, sternum 8; 10-12, genital capsule, lateral, ventral, and dorsal views.
Scale line = 1.00 mm.

be those of E. lata (Fig. 16); since the mandible of E. lata
possesses a distinct preapical tooth (wholly lacking in the
figure), the mandible figured is not that of E. lata. Also, they
stated that the tooth of the pollex is absent (p. 477); it is
present as the preapical tooth.

Hosts for Ericrocis are not known but, based on distri-

bution, will almost certainly prove to be species of Centris,
subgenus Paracentris, and possibly subgenus Acritocentris.

INCLUDED NAMES

arizonensis Baker, 1 906 ( Ericrocis )
lata (Cresson, 1878) (? Crocisa )

12 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

Figures 13-17. Acanthopus palmatus, male. 13, sternum 7; 14, sternum 8; 15-17, genital capsule, lateral, ventral, and dorsal views. Scale
line = 1 .00 mm.

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 13

Figures 18-22. Hopliphora velutina, male. 18, sternum 7; 19, sternum 8; 20-22, genital capsule, lateral, ventral, and dorsal views. Scale
line = 1.00 mm.

melectoides Baker, 1906 ( Ericrocis )
pintada Snelling and Zavortink, 1985 ( Ericrocis )
rossi Linsley, 1939 ( Ericrocis )
rugosa Fox, 1893 (Ericrocis)

Hopliphora Group

The two genera that comprise this group are characterized
by the lack of tergal patches or bands of appressed pale hairs,
the presence of a stemopleural ridge (weak in some Hopli-

phora), and the relatively long paraglossa (at least two-thirds
as long as the prementum).

(A) Inner eye margins slightly convergent above. (B) Ocel-
locular distance equal to diameter of anterior ocellus. (C)
Mandible with or without preapical tooth. (D) Maxillary pal-
pus absent or three-segmented. (E) Hypostomal carina low,
uniform. (F) Malus of protibia slender, shorter than velum,
without teeth. (G) Mesobasitarsus compressed and carinate
along posterior margin; distal process present, or not com-
pressed and without distal process. (H) Meso- and meta-

14 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

distitarsi without lateral patches of short, flattened setae. (I)
Metatarsus with or without posterior fringe. (J) Marginal cell
of forewing about 0.75-1.8 times distance from its apex to
wing tip; third submarginal cell usually more or less trian-
gular. (K) Juncture of basal and discal faces of first tergum
rounded. (L) Male seventh tergum bidentate at apex. (M)
Male gonostylus with or without dorsal lobe.

The group is represented by the two South American gen-
era Hopliphora and Acanthopus.

Genus Hopliphora Lepeletier

Figures 18-27, 61

Hopliphora Lepeletier, 1841:458. Type species: Mesocheira
velutina Lepeletier and Serville, 1825 (monobasic).
Eurytis F. Smith, 1854:279. Type species: Eurytis funereus
F. Smith, 1854 (monobasic).

Oxynedys Schrottky, 1902:491. Type species: ( Oxynedys be-
roni Schrottky, 1902) = Mesocheira velutina Lepeletier
and Serville, 1825 (monobasic and original designation).
Cyphomelissa Schrottky, 1902:493. Type species: (Cypho-
melissa pernigra Schrottky, 1902) = Melissa diabolica
Friese, 1 900 (monobasic and original designation). NEW
SYNONYMY.

Oxynedis: Moure, 1946:18, 27, 31 (lapsus).

DIAGNOSIS

Third submarginal cell receiving second recurrent vein only.

DESCRIPTION

(1) Head distinctly broader than long; inner eye margins
essentially straight, slightly convergent above; occipital mar-
gin distinctly elevated above ocelli. (2) First segment of max-
illary palpus distinctly shorter than second or third. (3) La-
brum with transverse preapical ridge; apical margin convex
to concave. (4) Interantennal distance distinctly greater than
antennal socket diameter; antennocular distance distinctly
less than antennal socket diameter. (5) Ocelloecipital distance
greater than diameter of anterior ocellus. (6) Occipital margin
abruptly rounded. (7) Antenna short in both sexes; minimum
length of first flagellar segment less than width at apex and
less, usually distinctly so, than length of second segment on
same side.

(8) Pronotum not carinate between collar and posterior
lobe. (9) Midline of mesoscutum broadly and deeply im-
pressed on anterior three-fourths, less strongly so toward
posterior margin; parapsidal lines not impressed. ( 1 0) Dorsal
face of scutellum broadly impressed along midline and with
a pair of suberect to erect blunt to subacute tubercles. (11)
Anterior mesepistemal carina and acetabular carina absent;
stemopleural ridge present, but often weak. (12) Supraspi-
racular ridge strong, terminating in stout, tooth-like process
well above, and slightly behind, spiracle.

(13) Tegula (Fig. 63) oval, outer margin evenly curved.

(14) Second submarginal cell, on M, as wide as, or wider
than, first; third submarginal cell not at all narrowed, or
greatly narrowed, anteriorly; 1st m-cu distinctly basad of 1st
r-m; 2nd m-cu at, or a little beyond, middle of third sub-
marginal cell; marginal cell 0.75 times as long as distance
from its apex to wing tip.

( 1 5) Mesotibial spur robust and very broad at apex (slender
and parallel-sided in one species), outer distal tooth often
indistinguishable from prominent intercalary teeth. (16)
Metatrochanter rounded beneath; metatibia with two normal
spurs; metadistitarsus 2. 5-3. 5 times longer than wide.

(17) Female sixth tergum with well-defined, sharply mar-
gir.ate pygidial plate, apex acute and reflexed, or narrowly
rounded and flat. (18) Male fourth sternum with distal mar-
gins straight or broadly emarginate, with distal fringe of more
or less prostrate long, dark, plumose hairs. (19) Male fifth
sternum either fully exposed and with straight apical margin
or hidden and with deeply emarginate margin, with or with-
out distal fringe. (20) Male seventh sternum with margin
angularly (Fig. 18) or broadly (Fig. 23) produced. (21) Male
eighth sternum variously produced. (22) Male gonostylus
short, thick, blunt; dorsal lobe present (Fig. 22) or absent
(Fig. 27); inner apical sclerotized portion of gonocoxite dis-
tinct. (23) Penis valve evenly curved ventrad; basolateral lobe
obsolete (Figs. 22-27).

DISCUSSION

Those species in which the third submarginal cell is trian-
gular, or even petiolate, have traditionally been placed in the
separate genus Cyphomelissa. However, species assigned to
Hopliphora have the third submarginal cell so strongly nar-
rowed anteriorly that the difference becomes one of degree
only. It is not enough, in our opinion, to place these two
groups of species in separate genera.

Superficially, some species of Mesonychium look much
like small Hopliphora and were included in Cyphomelissa
by Schrottky (1902). The following names are applicable to
Hopliphora as we understand the genus.

INCLUDED NAMES

beroni (Schrottky, 1 902) (Oxynedys)
commata (Moure, 1958) ( Cyphomelissa )
diabolica (Friese, 1900) ( Melissa )
funereus (F. Smith, 1854) ( Eurytis )
iheringi (Schrottky, 1 902) (. Acanthopus )
magnifiea (Moure, 1958) ( Cyphomelissa )
pernigra (Schrottky, 1 902) (Cyphomelissa)
superba (Ducke, 1 902) ( Melissa )
velutina (Lepeletier and Serville, 1825) (Mesocheira)

Genus Acanthopus Klug

Figures 13-17, 62, 70, 71

Acanthopus Klug, 1807:199, 226. Type species: (Apis splen-
dida Fabricius, 1793) = Apis palmata Olivier, 1789
(monobasic).

DIAGNOSIS

Third submarginal cell receiving both recurrent veins.
DESCRIPTION

(I) Head distinctly broader than long; inner eye margins
essentially straight, distinctly convergent above; occipital

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 15

Figures 23-27. Hopliphora superba, male. 23, sternum 7; 24, sternum 8; 25-27, genital capsule, lateral, ventral, and dorsal view. Scale line =
1.00 mm.

margin broadly convex and distinctly elevated above ocelli.
(2) Maxillary palpus absent. (3) Labrum with narrow, short
transverse preapical ridge; distal margin weakly, broadly con-
vex. (4) Interantennal distance less than antennal socket di-
ameter. (5) Ocelloccipital distance about three times diam-
eter of anterior ocellus. (6) Occipital margin abruptly rounded.
(7) Antenna short in both sexes; minimum length of first
flagellar segment greater than apical width and greater than
length of second or third segments on same side.

(8) Pronotum not carinate between collar and posterior
lobe. (9) Midline of mesoscutum deeply impressed for entire
length; parapsidal lines not impressed. (10) Dorsal face of
scutellum deeply impressed in middle, with a pair of erect,
conical tubercles. (1 1) Anterior mesepistemal carina present;
acetabular carina obsolete in middle; stemopleural ridge
present. (12) Supraspiracular ridge weak, evanescent distad.

( 1 3) Tegula (Fig. 62) elliptical, outer margin evenly curved,
posterior margin oblique. ( 1 4) Second submarginal cell about

16 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

Figures 28-33. Mesonychium coerulescens. 28, metadistitarsus, lateral view; 29, male sternum 7; 30, male sternum 8; 31-33, male genital
capsule, lateral, ventral, and dorsal views. Scale line = 1.00 mm (Figs. 29-33 only).

as broad as long; third submarginal cell on M longer than
first or second, receiving 1 st and 2nd m-cu, latter near mid-
length; marginal cell 1.8 times as long as distance from its
apex to wing tip.

( 1 5) Mesotibial spur wide, outer spine much longer than
inner, surface between strongly oblique and with three long,
widely spaced intercalary teeth. ( 1 6) Metatrochanter rounded
beneath; metatibial spurs normal, outer spur strongly curved
at tip; metadistitarsus about six times longer than wide.

( 1 7) Female sixth tergum with short, subtruncate pygidial
plate. (18) Male fourth sternum deeply, subangularly incised
and with a prostrate fringe of long hairs. (19) Male fifth
sternum largely hidden, posterior margin deeply incised and
with distal fringe of prostrate hairs. (20) Male seventh ster-
num (Fig. 1 3) with posterior margin broadly and somewhat
irregularly produced, apically acuminate. (21) Male eighth
sternum (Fig. 1 4) moderately produced, apex weakly emar-
ginate. (22) Male gonostylus short, broad, thickly digitiform

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 17

18 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

Table 3. Known or suspected hosts of Ctenioschelini.

Parasitoid

Host

Authority

Mesoplia dugesi

Centris anomala Snelling

Snelling, 1984'

Mesoplia rufipes

Centris carrikeri Cockerell

Rozen, 1969

Centris inermis Friese

Coville et al., 1983

(as C. segregala Crawford)
Epicharis albofasciata F. Smith

Rozen, 1969

Mesonychium gayi

Centris chilensis Spinola

Wagenknecht, 1974

Centris cineraria F. Smith

Wagenknecht, 1974

Centris nigerrima Spinola

Wagenknecht, 1974

Centris rhodophthalma Perez

Wagenknecht, 1974

Mesonychium jenseni

Centris autrani Vachal

Wagenknecht, 1974

Mesonychium wagenknechti

Centris chilensis Spinola

Wagenknecht, 1974

Centris rhodophthalma Perez

Wagenknecht, 1974

Abromelissa lendliana

Centris cineraria F. Smith

Wagenknecht, 1974

Centris nigerrima Spinola

Wagenknecht, 1974

Centris orellanai Ruiz

Wagenknecht, 1974

Centris rhodophthalma Perez

Wagenknecht, 1974

Aglaomelissa duckei

Centris carrikeri Cockerell

NEW RECORD2

Acanthopus palmatus

Ptilotopus derasus (Lepeletier)

Rozen, 1969

(as A. splendidus urichi)

1 Suspected host— observed at nest site.

2 Suspected host— observed at nest site (J.G. Rozen, Jr., personal communication).

in profile; dorsal lobe long, slender; inner apical sclerotization
of gonocoxite small but distinct. (23) Penis valve abruptly
deflected ventrad; basolateral lobe absent (Fig. 1 7).

DISCUSSION

The unusual wing venation (elongate marginal cell and sec-
ond submarginal cell receiving both recurrent veins) and the
conspicuous peculiar fringes on the metatarsi will immedi-
ately separate Acanthopus from other genera of ericrocines.

There is apparently but a single species, ranging from Trin-
idad and the Guianas to Brazil. The one known host is a
species of Ptilotopus (Table 3).

INCLUDED NAMES

excellerts Schrottky, 1902 (. Acanthopus )
jheringi Friese, 1 904 ( Acanthopus )
palmata (Olivier, 1789) (Apis)
splendida (Fabricius, 1793) (Apis)
urichi Cockerell, 1926 (Acanthopus)

Mesoplia Group

Diagnostic for this group are the elongate segments of the
maxillary palpus, the male lacks felt-like pads of pubescence
along the apical margins of metasomal sterna 4 and 5 (shared

with the Mesonychium group), and male sternum 5 is broadly
emarginate and largely concealed under sternum 4. The inner
eye margins are approximately parallel, rather than divergent
above as in the Mesonychium group.

(A) Inner eye margins weakly to distinctly divergent above.
(B) Ocellocular distance less than, equal to, or greater than
diameter of anterior ocellus. (C) Mandible with adnate pre-
apical tooth. (D) Maxillary palpus three- or four-segmented.
(E) Hypostomal carina moderately high, lamelliform. (F)
Malus of protibia short to long, usually with one or two teeth
along inner margin. (G) Mesobasitarsus compressed and car-
inate along posterior margin; distal process present. (H) Meso-
and metadistitarsi without lateral patches of short, flattened
setae. (I) Metatarsus without posterior fringe. (J) Marginal
cell of forewing 0.90-0.94 distance from its apex to wing tip.

(K) Juncture of basal and discal faces of first tergum rounded.

(L) Male seventh tergum bilobate at apex. (M) Male gono-
stylus with dorsal lobe.

Only the genus Mesoplia is included in this group which
ranges from the southwestern United States (Arizona) to Ar-
gentina.

Genus Mesoplia Lepeletier

Figures 1-12, 63, 64, 75

Mesoplia Lepeletier, 1841:457. Type species: Mesocheira
azurea Lepeletier and Serville, 1825 (monobasic).

Figures 34-38. Ericrocis lata, male. 34, sternum 7; 35, sternum 8; 36-38, genital capsule, lateral, ventral, and dorsal views. Scale line =

1 .00 mm.

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 19

Melissa F. Smith, 1854:279. Type species: Mesocheira azu-

rea Lepeletier and Serville, 1825 (designation of Sand-

house, 1943:570).

DIAGNOSIS

Same as description for Mesop/ia group.

DESCRIPTION

(1) Head distinctly broader than long; inner eye margins
essentially straight, subparallel to divergent above; occipital
margin nearly straight to low-convex, slightly, or not at all,
elevated above ocelli. (2) Maxillary palpus, except first seg-
ment, slender and elongate, segments more or less fused but
recognizable. (3) Labrum with transverse preapical ridge; api-
cal margin subtruncate. (4) Interantennal distance greater
than antennal socket diameter; antennocular distance less
than antennal socket diameter. (5) Ocelloccipital distance
equal to, or greater than, diameter of anterior ocellus. (6)
Occipital margin rounded. (7) Antenna short in both sexes;
minimum length of first flagellar segment less than apical
width and distinctly less than length of second segment on
same side.

(8) Pronotum with or without carina between collar and
front of posterior lobe. (9) Midline of mesoscutum impressed
for most of its length; parapsidal lines weakly, or not, im-
pressed. (10) Dorsal face of scutellum impressed along mid-
line, with suberect to erect mammiform tubercle on each
side. (11) Anterior mesepistemal carina, acetabular carina
and stemopleural ridge absent. (12) Supraspiracular ridge
weak, terminating well before spiracle.

(13) Tegula shape as in Figures 60 and 61, narrowed in
front, outer margin more or less sinuate. (14) First submar-
ginal cell, on M, as wide as, or wider than, second or third;
second and third submarginal cells narrowed anteriorly; 1st
m-cu interstitial with 1 st r-m; 2nd m-cu at, or a little distad
of, middle of third submarginal cell.

( 1 5) Mesotibial spur usually moderately broadened distad
and with one or more distinct intercalary teeth, but may be
parallel-sided for entire length. ( 1 6) Metatrochanter rounded
beneath; metatibial spurs of female normal; of male, normal
or with two very short, stout spurs or with a single short,
stout spur; metadistitarsus two and one-half to three times
longer than wide.

(17) Female sixth tergum with well-defined pygidial plate,
apex narrowly to broadly rounded. (18) Male fourth sternum
weakly to strongly concave along apical margin, with or with-
out distal fringe of long, plumose hairs. (19) Male fifth ster-
num hidden, distal margin deeply incurved and usually with
distal fringe of long, plumose hairs. (20) Male seventh ster-
num either (a) quadrately produced distad and with dense
patch of dark, bristle-like setae (Fig. 3) or (b) with posterior
margin more or less triangular, apex acute, truncate or bi-
lobed, and with scattered normal setae (Fig. 8). (21) Male
eighth sternum with small apical process which may be acute,
truncate or bilobate (Figs. 4, 9). (22) Male gonostylus short,
broad, thick and deflected ventrad (Figs. 5-7) or short, broad,
thin and curved mesad above the inner apical sclerotization

of the gonocoxite (Figs. 10-12). (23) Penis valves evenly
curved, or abruptly deflected, ventrad; basolateral lobe dis-
tinct.

DISCUSSION

Mesop/ia includes a dozen or so species ranging from south-
ern Arizona in the United States southward to northern Ar-
gentina. The genus is apparently absent from Chile and most
of the species occur in the Amazonian region of South Amer-
ica.

We have herein divided Mesoplia into two subgenera. The
males of the two subgenera are especially different in details
of the genitalia and associated sterna and these segregates
may be better regarded as distinct genera. However, the mag-
nitude of differences in the males is not reflected in the fe-
males. Until all of the species of Mesoplia in the broad sense
can be critically examined, we believe that generic separation
of the two segregates recognized here would be premature.

Known and suspected hosts are all species of Centris and
Epicharis (Table 3).

Subgenus Mesoplia

Figures 1-7, 60

DIAGNOSIS

MALE. With a single metatibial spur or with two very
short, stout spurs, the longer not extending beyond basal one-
third of metabasitarsus; metafemur often with robust basal
tooth-like projection ventrally; metatibia with, inner, distal
patch of black hairs. Female: pygidium broad, densely cov-
ered with appressed iridescent scales; dorsal surface of meta-
coxa angulate or carinate at juncture with anterior face.

DESCRIPTION

(24) Ocellocular distance equal to, or (usually) greater than,
diameter of anterior ocellus. (25) Pronotum with or without
low carina from collar to front of posterior lobe. (26) Dorsal
face of metacoxa sharply subangulate or carinate where it
meets anterior and posterior faces. (27) Metafemur of male
with or without stout, tooth-like basoventral projection. (28)
Metatibia of male with inner, distal hair patch and with one
or two apical spurs; if with two apical spurs, both short and
stout, neither extending beyond basal one-third of metaba-
sitarsus. (29) Tegula shape as in Figure 60. (30) Female py-
gidium broad, apex broadly rounded, disc covered with ap-
pressed metallic scales. (31) Male seventh sternum (Fig. 3)
quadrately produced and with distal patch of short, dense,
stout, black setae. (32) Male gonostylus short, broad, thick,
deflected ventrad (Fig. 5); dorsal lobe short, broad. (33) Ba-
solateral process of penis valve relatively stout (Fig. 7).

DISCUSSION

This subgenus, with species ranging from the southwestern
United States to Argentina, includes two distinctive species
groups. In both sexes of the M. azurea group ( sensu Moure,
1960a, b) there is a low carina extending laterad from the

20 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

pronotal collar to the front of the posterior lobe. In the males,
the metafemur has a stout, basal projection on the ventral
surface and there is a single metatibia! spur.

The M. bifrons group ( sensu Moure, 1960a, b) includes
species that do not possess a carina between the pronotal
collar and the posterior lobe. Males do not have a basoventral
projection on the metafemur and there are two short, stout
metatibial spurs.

In the following list, names marked by an asterisk (*) be-
long to the M. azurea group and those marked with a dagger
(f) belong to the M. bifrons group; group placement is un-
certain for those names that are unmarked.

INCLUDED NAMES

*azurea (Lepeletier and Serville, 1825) ( Mesocheira )
f bifrons (Fabricius, 1 804) (. Melecta )
ckalybea (Friese, 1912) ( Melissa )
chiruana (Holmberg, 1885) (Melissa)

*dugesi (Cockerell, 1917) ( Mesonychium )

*guaiemalensis Cockerell, 1912 ( Mesoplia )

*imperialis (Ashmead, 1 900) ( Melissa )

f imperialis (Friese, 1912) ( Melissa ) PREOCCUPIED

f imperatrix (Friese, 1913) ( Melissa )

t insignis (F. Smith, 1879) ( Melissa )

itaitubina (Ducke, 1902) ( Melissa )

maculata (Friese, 1900) ( Melissa )

ornata (Spinola, 1841) ( Mesocheira )

f pilicrus (Friese, 1902) ( Melissa )

pretiosa (Friese, 1912) ( Melissa )

f regalis (F. Smith, 1854) ( Melissa )

*rufipes (Perty, 1833) ( Crocisa )
simillima Schrottky, 1 920 (Mesoplia)

Eumelissa, new subgenus

Figures 8-12, 61

DIAGNOSIS

MALE. Metatibial spurs normal, inner spur extending to,
or beyond, midlength of metabasitarsus; metafemur without
basal tooth; metatibia without inner, distal seta patch. Fe-
male: pygidium narrow', shiny, surface bare; metacoxa round-
ed above, neither carinate nor angulate at juncture with an-
terior face.

DESCRIPTION

(24) Ocellocular distance equal to, or less than, diameter of
anterior ocellus. (25) Pronotum without carina between col-
lar and front of posterior lobe. (26) Metacoxa abruptly round-
ed between dorsal and lateral faces, not angulate or subcar-
inate. (27) Metafemur of male without basal tooth or
projection. (28) Metatibia of male without inner, distal patch
of setae and with two normal apical spurs, the longer ex-
tending to, or beyond, midlength of metabasitarsus. (29) T eg-
ula (Fig. 61) elliptical, narrowed behind. (30) Female pygid-
ium completely marginate, narrow, apex narrowly rounded,
disc bare and shiny. (31) Female sixth sternum weakly car-
inate along midline. (32) Male seventh sternum (Fig. 8) with

apical margin somewhat triangularly produced in middle,
apex acute, truncate or bilobate, with a few short, simple
setae. (33) Gonostylus of male short, broad, curved mesad
over inner plate (Fig. 11); dorsal lobe elongate. (34) Baso-
lateral process of penis valve relatively slender (Fig. 12)

TYPE SPECIES

Melissa decorata F. Smith, 1854.

ETYMOLOGY

Greek prefix eu- (beautiful) plus melissa (bee).

DISCUSSION

Males of this subgenus are easily recognized by the unmod-
ified metafemur and metatibia, as well as by the genitalia
and associated structures. Both sexes share the unmodified
metacoxa, the dorsal surface of which is distinctly rounded
into the anterior and posterior surfaces, rather than carinate
or sharply subangulate as in species of Mesoplia s. str. Fe-
males otherwise are very similar to those of the nominate
subgenus but differ in the narrow, shiny pygidial plate which
is devoid of the dense covering of metallic scales character-
istic of that subgenus.

The included species are all South American, but we have
seen both sexes of a possibly undescribed species from Costa
Rica. Moure (1960b) included Melissa duckei Friese with
this group of species, but in our opinion this species is not
congeneric and is the type species for the genus Aglaomelissa,
described below.

INCLUDED NAMES

albogntta (Ducke, 1905) (Melissa)
albopunctata Moure, 1967 (Mesoplia)
decorata (F. Smith, 1854) (Melissa)
friesei (Ducke, 1 902) (Melissa)
guedesii (Ducke, 1 902) ( Mesocheira )

Mesonychium Group

Diagnostic characteristics of this group are inner eye margins
divergent above (shared with Ericrocis group); strigilar malus
without teeth (shared with Aglaomelissa in the Ctenioschelus
group); the mesocoxa is less than 1.35 times the distance
from the summit of the coxa to the base of the hind wing;
metasomal sternum 3 of the male is without a dense lateral
patch of white hair; and, male sterna 4 and 5 are without
felt-like pads of pubescence along the apical margins. Me-
tallic, scale-like hairs are present in Mesonychium, but absent
in Abromelissa, in which the metasomal integument is me-
tallic bluish, a unique apomorphy within the Ericrocini.

(A) Inner eye margins divergent above, often strongly so.
(B) Ocellocular distance greater than diameter of anterior
ocellus. (C) Mandible with preapical tooth. (D) Maxillary
palpus one- or two-segmented. (E) Hypostomal carina low,
uniform. (F) Malus of protibia short, one-half or less length
of velum, inner margin simple. (G) Mesobasitarsus usually

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 21

compressed and more or less sharply carinate along posterior
margin; distal process usually present. (H) Meso- and meta-
distitarsi with lateral patch of short, flattened setae on each
side (Fig. 28) (except Abromelissa and a few Mesonychium).
(I) Metatarsus without posterior fringe. (J) Marginal cell of
forewing 0.60-0.94 times distance from its apex to wing tip.

(K) Juncture of basal and discal faces of first tergum rounded.

(L) Male seventh tergum bilobate at apex or with two stout
teeth. (M) Gonostylus without dorsal lobe.

The two included genera, Mesonychium and Abromelissa
are confined to South America.

Genus Mesonychium Lepeletier and Serville

Figures 28-33, 65

Mesonychium Lepeletier and Serville, 1 825: 1 07. Type species;
Mesonychium coeru/escens Lepeletier and Serville, 1825
(monobasic).

Epiclopus Spinola, 1851:183. Type species: Epiclopus gayi
Spinola, 1851 (monobasic).

DIAGNOSIS

Abdomen with appressed metallic scales or hairs; labrum
with transverse preapical ridge which may be divided in
middle; scutellum usually bituberculate.

DESCRIPTION

( 1 ) Head much broader than long; inner eye margins straight,
moderately to strongly divergent above; occipital margin lit-
tle, if any, elevated above ocelli. (2) Maxillary palpus con-
sisting of a single short, spindle-shaped segment. (3) Apical
margin of labrum subtruncate; preapical transverse ridge
(sometimes interrupted in middle) present. (4) Interantennal
distance greater than antennal socket diameter; antennocular
distance greater than antennal socket diameter. (5) Ocelloc-
cipital distance greater than diameter of anterior ocellus. (6)
Occipital margin abruptly rounded to subangulate. (7) An-
tenna short in both sexes; minimum length of first flagellar
segment distinctly less than, to about equal to, apical width
and much shorter than, to as long as, length of second seg-
ment on same side.

(8) Pronotum not carinate between collar and posterior
lobe. (9) Midline of mesoscutum impressed for nearly its
entire length; parapsidal lines weakly impressed, if at all. (10)
Dorsal face of scutellum broadly, often weakly, impressed
along midline; with a pair of erect, mammiform tubercles or
(one species) no tubercles. (11) Anterior mesepistemal carina,
acetabular carina, and stemopleural ridge absent. (12) Su-
praspiracular ridge weak to moderately strong, terminating
in a stout tooth or projection above spiracle. ( 1 3) Tegula (Fig.
64) elliptical, outer margin evenly curved. (14) Second sub-
marginal cell, on M, wider than, or equal to, first and wider
than third; third submarginal cell narrowed anteriorly, some-
times triangular or petiolate; 1st m-cu basad of, or interstitial
with, 1st r-m; 2nd m-cu distinctly basad of 2nd r-m.

(15) Mesotibial spur long, slender, parallel-sided, outer
distal tooth obsolete. (16) Metatrochanter rounded beneath;
metatibial spurs normal; metadistitarsus less than three times
longer than wide.

(17) Female sixth tergum with short, distinct pygidial plate,
apex narrowly rounded to subtruncate. (18) Male fourth ster-
num usually concealed under third, its apical margin broadly,
deeply incurved. (19) Male fifth sternum exposed, apical mar-
gin broadly, shallowly incurved or subtruncate. (20) Male
seventh sternum (Fig. 29) more or less produced in middle,
subtruncate to bilobate at apex. (21) Male eighth sternum
(Fig. 30) with apical margin little produced. (22) Male gono-
stylus short and broad in dorsal view, stoutly digitiform in
lateral view; dorsal lobe absent; inner apical sclerotization
of gonocoxite usually well defined. (23) Penis valve weakly
curved ventrad; basolateral lobe prominent (Fig. 33).

DISCUSSION

Mesonychium has never been adequately distinguished from
Mesoplia by previous workers. Although species in the two
genera are often somewhat similar in appearance, the two
are distinct. Most species of Mesonychium, for example, pos-
sess very distinct patches of flattened setae on the meso- and
metadistitarsi, which are lacking in Mesoplia. Male gonostyli
lack dorsal lobes in species of Mesonychium and in both
sexes of this genus the inner eye margins are moderately to
strongly divergent above. Mesonychium is similar to the
Nearctic genus Ericrocis, from which it is easily separated
by the presence of metallic blue or green hairs and/or scales
on various areas of the body.

Mesonychium appears to be an exclusively South Ameri-
can genus with species in Peru and Brazil south to Argentina
and Chile. Within the genus there are two principal groups.
The first of these includes the generotype and a few other
species in which the meso- and metadistitarsi bear a cluster
of short, flattened setae on each side and the pubescence of
the head and body are generally dark. In a few species in this
group the pubescence of the thoracic dorsum is very short
and sparse. Species assigned to the M. coerulescens group
include: M. asteria (F. Smith), M. garleppi (Schrottky), M.
jenseni (Friese), M. littoreum Moure, M. viridescens (Friese),
and M. viridis (Friese).

A second group of species lacks the distitarsal setae and
in most the thorax and first two abdominal segments are
densely clothed with long, erect white hairs; at least one
species is dark haired. The species of this second group in-
clude those previously assigned to Epiclopus, but this group
is so diverse in its morphological features that use of that
name, even as a subgenus, would not be appropriate at this
time. The following species belong to this group: M. gayi
(Spinola), M. wagenknechti Ruiz, and an undescribed Pe-
ruvian species.

Although M. lendlianum (Friese) has traditionally been
associated with such other Chilean species as M. gayi, which
it resembles, it is anomalous in Mesonychium, and is the
sole representative of the new genus Abromelissa, described
below.

INCLUDED NAMES

albescens (Friese, 1921) (Melissa)
andina (Friese, 1925) ( Melissa )
asteria (F. Smith, 1854) ( Mesocheira )

22 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

coerulescens Lepeletier and Servilie, 1825 (. Mesonychium )

chilensis (F. Smith, 1854) ( Melecta )

garleppi (Schrottky, 1910) ( Cyphomelissa )

gayi (Spinola, 1851) ( Epiclopus )

jenseni (Friese, 1 906) ( Melissa )

littoreum Moure, 1944 ( Mesonychium )

porteri (Herbst, 1917) ( Melissa )

violacea (Friese, 1 900) (Melissa)

viridescens (Friese, 1930) ( Melissa )

viridis (Friese, 1 900) ( Melissa )

wagenknechti Ruiz, 1938 [ Mesonychium ( Epiclopus )]

Abromelhsa, new genus

Figures 39-43, 66

DIAGNOSIS

Separable from all other ericrocine genera by the metallic
blue integument of the abdominal terga, without appressed
metallic-reflective scales or hairs; further separable from Me-
sonychium and Ericrocis by the presence of a long cylindrical
dorsal lobe on the male gonostylus, and in both sexes by the
broader than long abdomen.

DESCRIPTION

(1) Head much broader than long; inner eye margin straight,
moderately divergent above; ocelli on top of preocciput. (2)
Maxillary palp three-segmented, first segment very short,
second more than twice longer than first and about 1.5 times
longer than third. (3) Apical margin of labrum subtruncate;
preapical transverse ridge present and entire. (4) Interanten-
nal distance greater than antennal socket diameter; anten-
nocular distance greater than antennal socket diameter. (5)
Ocelloccipital distance greater than diameter of anterior ocel-
lus. (6) Occipital margin abruptly rounded. (7) Antenna short
in both sexes; minimum length of first flagellar segment great-
er than apical width and greater than length of second seg-
ment on same side.

(8) Pronotum not carinate between collar and posterior
lobe. (9) Midline of mesoscutum impressed almost to pos-
terior margin; parapsidal lines not impressed. (10) Dorsal
face of scuteilum barely impressed along midline and without
mammiform tubercles, but with a pair of sublateral, short,
erect spines at margin of declivity. (II) Anterior mesepi-
stemal carina, acetabular carina and stemopleural ridge ab-
sent. (12) Supraspiracular ridge strong, ending in a stout,
blunt projection above spiracle.

(13) Tegula pyriform (Fig. 66). (14) Second submarginal
cell, on M, wider than either first or third; third submarginal
cell narrower on Rs than on M; 1st m-cu interstitial with 1st
r-m; 2nd m-cu much basad of 2nd r-m.

(15) Mesotibial spur long, slender, parallel-sided, outer
distal tooth obsolete. (16) Metatrochanter rounded beneath;
metatibial spurs normal; metadistitarsus less than three times
longer than wide and without lateral patch of short, flattened
setae.

(17) Female sixth tergum with short, distinct pygidial plate,
apex narrowly rounded. (18) Male fourth sternum exposed,
its apical margin transverse and with a dense fringe of de-

cumbent, long, dark hairs. (19) Male fifth sternum normally
visible only at extreme sides, its apical margin broadly and
shallowly incurved and partially concealed under fourth. (20)
Male seventh sternum (Fig. 39) with well-developed distal
lobe, weakly trilobate. (21) Male eighth sternum (Fig. 40)
with apical margin moderately produced. (22) Male gono-
stylus (Fig. 43) with dorsal, elongate, cylindrical lobe; inner
apical sclerotization of gonocoxite short, broad and concave
on dorsal face (Fig. 42), stout and subtruncate in lateral view
(Fig. 41). (23) Penis valve strongly curved ventrad; basolat-
eral lobe prominent.

TYPE SPECIES

Melissa (Epiclopus) lendliana Friese, 1910.

ETYMOLOGY

Combines Greek habros (graceful or pretty) with melissa
(bee).

DISCUSSION

This genus includes only the type species, found in Argentina
(Provinces of Neuquen and Valdivia) and Chile (Provinces
of Aconcagua, Coquimbo, Valparaiso, Santiago, Nuble, Cu-
rico, and Aisen). Melissa friesei Herbst, 1918 (not M. friesei
Ducke, 1902) and Mesonychium frieseanum Ruiz, 1938, are
synonyms of A. lendliana (NEW SYNONYMIES). The one
species of Abromelissa superficially resembles two Chilean
species of Mesonychium, M. gayi and M. wagenknechti, since
the hairs of the thorax and first tergum are long, erect and
pale, and there are no short, appressed, metallic-reflective
scales on the body. From both of these, however, it differs
in the dark metallic blue color of the tergal integument, a
feature which appears to be unique within the tribe.

In addition to the characteristics described above, there
are a few other features of Abromelissa that are distinctive
within the Mesonychium group. The surface of the mesoscu-
tum is smooth and shiny between subcontiguous coarse
punctures and sparse giant punctures, and the scuteilum is
coarsely rugosopunctate and foveolate on the dorsal face.
Although the mesobasitarsus is without a ridge along its pos-
terior margin, an inconspicuous distal process is present. The
male pygidial plate is narrowly translucent at its apex and is
weakly bilobate.

The known hosts are all species of Centris in the subgenera
Paracentris and Wagenknechtia (Table 3).

Ctenioschelus Group

Several unique apomorphies define the Ctenioschelus group:
the preapical tooth is lost; the forewing tips are infuscated;
metasomal tergum 7 is subangulate in profile (Fig. 76). Arolia
are present (except in Mesocheira) and an anterior mesepi-
stemal carina is present (except in Ctenioschelus).

(A) Inner eye margins subparallel or weakly convergent
above. (B) Ocellocular distance less than diameter of anterior
ocellus. (C) Mandible without preapical tooth, sometimes
fused with stipes. (D) Maxillary palpus consisting of one short
segment. (E) Hypostomal carina high, lamelliform. (F) Malus

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 23

Figures 39-43. Abromelissa lendlianum, male. 39, sternum 7; 40, sternum 8; 41-43, genital capsule, lateral, ventral, and dorsal views.

Scale line = 1.00 mm.

24 Contributions in Science, Number 369

Snell ing and Brooks: The Tribe Ericrocini

of protibia short (0.25 or less length of velum), stout, simple.
(G) Mesobasitarsus neither compressed nor carinate along
posterior margin; distal process absent. (H) Meso- and meta-
distitarsi without lateral patches of short, flattened setae. (I)
Metatarsus without posterior fringe. (J) Marginal cell of fore-
wing 1.0-1. 2 times distance from its apex to wing tip. (K)
Juncture of basal and discal faces of first tergum slightly
elevated and subangulate. (L) Male seventh tergum bidentate,
teeth well separated. (M) Male gonostylus absent or, if pres-
ent, without dorsal lobe.

The three genera, Aglaomelissa , Ctenioschelus, and Me-
socheira occur in Central and South America.

Aglaomelissa , new genus

Figures 44-48, 67

DIAGNOSIS

Separable from Ctenioschelus and Mesocheira by the follow-
ing combination of characteristics: male antenna short; an-
terior mesepistemal carina present; scutellar prominences
mammiform; marginal cell of forewing without apical cloud.

DESCRIPTION

( 1 ) Head a little broader than long; inner eye margins straight,
slightly convergent above; occipital margin elevated above
ocelli. (2) Maxillary palpal segment short, broad, lightly scler-
otized and fused to stipes. (3) Labrum without preapical
tubercle or transverse ridge; apical margin slightly produced
and truncate. (4) Interantennal distance slightly greater than
antennal socket diameter; antennocular distance about equal
to one-half antennal socket diameter. (5) Ocelloccipital dis-
tance about two times diameter of anterior ocellus. (6) Oc-
cipital margin subcarinate. (7) Antenna short in both sexes;
minimum length of first flagellar segment less than greatest
width and distinctly less than length of second segment on
same side.

(8) Pronotum not carinate between collar and posterior
lobe. (9) Midline of mesoscutum impressed almost to pos-
terior margin; parapsidal lines not impressed. (10) Dorsal
face of scutellum broadly impressed in middle and with a
pair of suberect mammiform tubercles. (11) Anterior mes-
epistemal carina sharp, lamelliform and confluent with ace-
tabular carina, or nearly so; stemopleural ridge absent. ( 1 2)
Supraspiracular ridge evanescent where its distal portion turns
ventrad.

(13) Tegula (Fig. 67) elliptical, outer margin narrowed an-
teriorly. ( 1 4) Submarginal cells about equally long on M, third
much narrowed anteriorly; 1st m-cu interstitial with 1st r-m;
2nd m-cu distinctly basad of 2nd r-m; marginal cell about
as long as distance from apex to wing tip.

(15) Mesotibial spur stout, apex broad, with 1-3 elongate
intercalary teeth. (16) Metatrochanter compressed and sub-
angular below; outer metatibial spur short, stout, strongly
curved at apex; metadistitarsus about twice longer than wide.

(17) Female sixth tergum with well-defined, narrow py-

gidial plate, apex subacute. ( 1 8) Male fourth sternum broadly
incurved and with distal fringe of long, plumose, prostrate
hairs. (19) Male fifth sternum more shallowly incurved and
with shorter distal fringe. (20) Male seventh sternum (Fig.
44) with median lobe on apical margin. (21) Male eighth
sternum (Fig. 45) with apical margin produced, emarginate,
and narrowly truncate in middle. (22) Male gonostylus absent
(Figs. 46, 47) inner apical sclerotization of gonocoxite dis-
tinct. (23) Penis valve evenly curved ventrad; basolateral lobe
absent.

TYPE SPECIES

Melissa ( Mesocheira ) duckei Friese, 1906.

ETYMOLOGY

Greek, aglaos (splendid or beautiful) plus melissa (bee, also
an old generic name in this tribe).

DISCUSSION

Although this genus is related to Ctenioschelus and Meso-
cheira, it is easily separated from both. Males of Ctenio-
schelus have extraordinarily long antennae, and both sexes
of that genus lack pronotal, anterior mesepistemal, and ace-
tabular carinae. In Mesocheira the processes of the dorsal
face of the scutellum are flattened and plate-like, extending
over the base of the abdomen.

The only species of Aglaomelissa is known from a few
specimens from Costa Rica, Panama, Trinidad, Colombia,
and Venezuela.

Although Moure (1960b) included A. duckei in his Me-
soplia decorata group, this bee is clearly not a Mesoplia.
Particularly indicative of its relationship to Ctenioschelus
and Mesocheira are the presence of the anterior mesepister-
nal carina and the subangulate profile of the first metasomal
tergum.

The suspected host is a species of Centris (Table 3).

Genus Ctenioschelus Romand

Figures 49-54, 68

Ctenioschelus Romand, 1840:336. Type species : Acanthopus
goryi Romand, 1 840 (monobasic).

Ischnocera Shuckard, 1840:166. No included species.
Melissoda Lepeletier, 1841:508. Type species: ( Melissoda la-
treillei Lepeletier, 1841) = Acanthopus gory>i Romand, 1 840
(monobasic).

DIAGNOSIS

Male immediately separable from all other ericrocine genera
by the greatly elongate flagellum reaching beyond apex of
abdomen. Additional features characteristic of both sexes:
no anterior mesepistemal carina; scutellar tubercles pros-
trate, mammiform, and extended over propodeum; meso-

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 25

44

Figures 44-48. Aglaomelissa duckei, male. 44, sternum 7; 45, sternum 8; 46-48, genital capsule, lateral, dorsal, and ventral views. Scale
line = 1.00 mm.

26 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

Figures 49-54. Ctenioschelus goryi. 49, female mesobasitarsus, pilosity omitted; 50, male sternum 7; 51, male sternum 8; 52-54, male genital
capsule, lateral, ventral, and dorsal views. Scale line = 1.00 mm.

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 27

Figures 55-59. Mesocheira bicolor, male. 55, sternum 7; 56, sternum 8; 57-59, genital capsule, lateral, ventral, and dorsal views. Scale line =
1.00 mm.

basitarsus without posterior carina or distal process; juncture
of basal and discal faces of first tergum elevated and angulate.

DESCRIPTION

(1) Head a little broader than long; inner eye margins essen-
tially straight, convergent above; occipital margin little ele-

vated above ocelli. (2) Maxillary palpal segment short, flat-
tened. (3) Labrum impressed along midline and with weak
transverse preapical ridge; apical margin broadly rounded,
subtruncate in middle. (4) Interantennal distance less than
antennal socket diameter; antennocular distance less than
antennal socket diameter. (5) Ocelloccipital distance more

28 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

63

Figures 60-69. Dorsal view of left tegula of: 60, Ericrocis lata ; 61, Hopliphora velutina ; 62, Acanthopus palmatus; 63, Mesoplia (M.) rufipes;
64, M. (Eurnelissa) decorata ; 65, Mesonychium coerulescens; 66, Abromelissa lendliana ; 67, Aglaomelissa ducket; 68, Ctenioschelus goryi; 69,
Mesocheira bicolor; scale line = 1 .0 mm. 70, diagrammatic mesopleuron, illustrating positions of: anterior mesepistemal carina (amc); acetabular
carina (acc); mesocoxa (cx 2); procoxa (cx 1); stemopleural ridge (spr). Precoxal depression stippled.

than twice diameter of anterior ocellus. (6) Occipital margin
subangulate. (7) Antenna short in female, flagellum longer
than body length in male; minimum length of female first
flagellar segment less than width and about one-half as long
as second segment; minimum length of male first flagellar

distinctly greater than width and less than one-third as long
as second on same side; middle segments of male flagellum
about seven times longer than wide.

(8) Pronotal collar prominent on each side, no carina be-
tween collar and posterior lobe. (9) Midline of mesoscutum

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 29

Figures 71-77. Acanthopus palmatus, female: 71-72, Posterior and side view of cardines (apices only), lorum, mentum, and basal portion
of prementum arranged in a single plane. Ericrocis lata, female: 73-74, posterior and side view of cardines (apices only), lorum, mentum, and
basal portion of prementum arranged in a single plane. 75, side view of metasoma, spiracles omitted. Mesoplia imperatrix, male: 76, posterior
view of cardines (apices only), lorum, mentum, and basal portion of prementum arranged in a single plane. Mesocheira bicolor, female: 77,
side view of metasoma, spiracles omitted, arrow indicates character 67. Abbreviations are A, basal apodeme of prementum; ACT, anterior
conjunctival thickening [=suspensory thickening of Winston (1979)]; C, cardo; K, notch of metasomal sternum 5; L, lorum; M, mentum; P,
prementum; Tl, first metasomal tergum. Dotted areas represent the membranous surface of the labiomaxillary tube extending toward its
attachment to the head.

not impressed; parapsidal lines weakly impressed. (10) Dor-
sal face of scutellum impressed along midline; lateral pro-
cesses nearly prostrate, dorsally convex, projecting over con-
cave vertical face, metanotum, and propodeum. (1 1) Anterior
mesepistemal carina, acetabular carina, and stemopleural
ridge absent. (12) Supraspiracular ridge prominent, shelf-like,
not curved ventrad behind spiracle.

( 1 3) Tegula subrectangular, outer margin sinuate (Fig. 68).

(14) First submarginal cell, on M, narrower than second or
third; third submarginal cell much narrowed anteriorly; 1st
m-cu interstitial with 1st r-m; 2nd m-cu distinctly basad of
2nd r-m; wing tip infuscated.

( 1 5) Mesotibial spur broad at apex, with two or three long

intercalary teeth. (16) Metatrochanter compressed and an-
gulate ventrally; metatibial spurs normal; metadistitarsus
about three times longer than wide.

(17) Female sixth tergum with short, poorly defined, nar-
rowly truncate pygidial plate. (18) Male fourth sternum
broadly emarginate, distal portion bare and transparent, but
largely hidden by long brush of hairs arising near midlength.
( 1 9) Male fifth sternum similar but transparent margin nar-
rower, more fully concealed by preapical fringe. (20) Male
seventh sternum (Fig. 50) rounded distad, subangulate in
middle. (21) Male eighth sternum (Fig. 51) produced along
apical margin, with small median emargination. (22) Male
gonostylus almost absent, situated beneath much enlarged

30 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

78

Figure 78. A. Cladogram showing the generic relationships of the Ericrocini and its sister group the Rhathymini which are shown derived
from the Centridini. Caenonomada has other apomorphies which have not been given. A bar indicates an apomorphy, an X a reversal.
Synapomorphies which root both trees are 20, 23, 24, 52. Synapomorphies of the centridine-rhathymine-ericrocine lineage (intemode 1-2)
are the derived characters of variables 17, 18, 20(2), 25, 26, 44, 47(2). When there is more than one derived character composing a variable
the relevant one is indicated in parentheses. Synapomorphies of the rhathymine-ericrocine lineage (intemode 2-3) are 2, 7, 14, 21(2), 29, 37,
39, 46, 49, 64. Synapomorphies of the Ericrocini (intemode 3-4) are 1, 10, 12, 19, 22, 27, 28, 31, 32, 35, 36, 45, 50, 53, 57, 58, 60, 61(2),
62. B. Alternative cladogram showing relationships of the tribes. Synapomorphies of rhathymine-centridine-ericrocine bees (intemode 5-6)
are 17, 18, 20(2), 25, 26, 47(2). Synapomorphies of Rhathymini are 2, 5, 7, 14(3), 21(3), 29, 37, 39, 45 (character is not constant), 46, 49,
58, 59, 64, 66. Synapomorphies of centridine-ericrocine bees (intemode 6-7) are 27 (character is not constant), 28, 3 1 (character is not constant),
44. Synapomorphies for Centridini (excluding Caenonomada) are 21 (reversion), 24 (reversion), 26(2). Synapomorphies of the Ericrocini are
1, 2, 7, 10, 12, 14, 19, 21(2), 22, 27-29, 31, 32, 35-37, 39, 43, 45-47, 49, 50, 57-60, 61(2), 62-64.

Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini 31

concave and densely setose inner apical sclerotization of
gonocoxite (Fig. 54). (23) Penis valve evenly curved ventrad;
basolateral lobe weak (Fig. 54).

DISCUSSION

The bizarrely slender and elongate antennae are immediately
distinctive for Ctenioschelus males. Females resemble those
of Mesoplia but are readily separable by the subangulate
profile of the first tergum. The one species is apparently not
common and ranges from Costa Rica to Brazil. Brazilian
specimens have their forewing tips more heavily infuscate
than do the Peruvian to Central American populations. Its
host is unknown.

INCLUDED NAMES

goryi (Romand, 1 840) ( Acanthopus )
latreillei (Lepeletier, 1841) ( Melissoda )

Genus Mesocheira Lepeletier and Serville

Figures 55-59, 69, 76

Mesocheira Lepeletier and Serville, 1825:106. Type species:
( Mesocheira bicolor Lepeletier and Serville, 1825) = Me-
lecta bicolor Fabricius, 1804 (designation of Taschenberg,
1883).

Mesochira Schulz, 1906:257 (lapsus).

DIAGNOSIS

Scutellar processes prostrate and shelf-like, extending over
base of abdomen; anterior mesepistemal carina present; mar-
ginal cell of forewing with apical cloud.

DESCRIPTION

(1) Head a little broader than long; inner eye margins essen-
tially straight, weakly convergent above; occipital margin not
elevated above ocelli. (2) Maxillary palpal segment short,
button-like. (3) Labrum impressed along midline, without
preapical ridge or tubercle; apical margin transverse or slight-
ly concave. (4) Interantennal distance about equal to antennal
socket diameter; antennocular distance less than antennal
socket diameter. (5) Ocelloccipital distance almost twice di-
ameter of anterior ocellus. (6) Occipital margin reflexed and
cariniform. (7) Antenna short in both sexes; minimum length
of first flagellar segment about one-half maximum width and
less than one-half length of second segment on same side.

(8) Side of pronotal collar raised and subcarinate, with high
lamelliform carina from collar to front of posterior lobe. (9)
Midline and parapsidal lines of mesoscutum weakly im-
pressed. (10) Dorsal face of scutellum narrowly impressed
along midline; lateral processes flat, shelf-like, extending over
base of abdomen. (11) Anterior mesepistemal carina lamel-
liform, confluent below with acetabular carina; stemopleural
ridge absent. (12) Supraspiracular ridge prominent, curved
ventrad for a short distance behind spiracle.

(13) Tegula subrectangular, outer margin sinuate (Fig. 69).

(14) First submarginal cell, on M, narrower than second or

third; third submarginal cell strongly narrowed anteriorly;
1st m-cu interstitial with 1st r-m; 2nd m-cu virtually inter-
stitial with 2nd r-m; wing clear, with apical infuscation in
marginal cell and at wing tip.

(15) Mesotibial spur stout, weakly broadened distad, with
one or two long, acute intercalary teeth. (16) Metatrochanter
compressed and subangulate beneath; metatibial spurs nor-
mal; metadistitarsus less than three times longer than wide.

(17) Female sixth tergum with well-defined, acute pygidial
plate. (18) Male fourth sternum with sharp, median, longi-
tudinal carina on basal one-half, distal margin nearly straight.
( 1 9) Male fifth sternum hidden under fourth, apical margin
deeply concave. (20) Male seventh sternum (Fig. 55) with
apical margin strongly produced, subtruncate and with me-
dian triangular projection. (2 1) Male eighth sternum (Fig. 56)
with medioapical portion produced, often irregular in shape.
(22) Male gonostylus almost absent, reduced to a mere slit
only visible from apicodorsal aspect (Fig. 59); inner apical
sclerotization of gonocoxite nearly absent. (23) Penis valves
evenly curved ventrad; basolateral lobe moderately pro-
duced.

DISCUSSION

The plate-like, posteriorly directed mesoscutellar processes
will separate both sexes of Mesocheira from all other ericro-
cine genera. There appears to be but a single species that
ranges from Mexico to Brazil and Paraguay. Although hosts
are unknown, the relatively small size suggests species of the
subgenera Hemisiella and Heterocentris of Centris. The se-
nior author has taken females at a nest site of C. ( Hemisiella )
nitida F. Smith in Mexico.

The figures of the labiomaxillary complex of the mouth-
parts, illustrated (fig. 44) by Winston (1979) as those of Me-
socheira bicolor, are not of that bee. They appear to be based
on Ericrocis lata.

INCLUDED NAMES

bicolor (Fabricius, 1 804) ( Melecta )
bilamellosa (Cockerell, 1949) ( Exaerete )
elizabethae Cockerell, 1910b ( Mesocheira )
melanura (Cockerell, 1949) ( Exaerete )
pulchella Holmberg, 1887 ( Mesocheira )
sericea Guerin-Meneville, 1846 ( Mesocheira )

ACKNOWLEDGMENTS

For the loan of specimens seen during the course of this
review we are very much indebted to the following colleagues:
G.C. Eickwort, Cornell University; the late P.D. Hurd, Jr.,
United States National Museum of Natural History; E.G.
Linsley, University of California, Berkeley; C.D. Michener,
University of Kansas; A.R. Moldenke, Oregon State Uni-
versity; F.D. Parker and T. Griswold, Utah State University;
W.J. Pulawski, California Academy of Sciences; J.G. Rozen,
Jr., and M. Favreau, American Museum of Natural History;
R.O. Schuster, University of California, Davis.

Our thanks to Dr. Charles D. Michener who first discov-

32 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

ered many of the variables used in the cladistic analysis and
for his valuable suggestions in the preparation of the clado-
gram. Joetta Weaver helped with the typing of the Cladistic
section.

This study was partially funded by National Science Foun-
dation grant DEB82- 12223 (C.D. Michener, principal in-
vestigator), which is gratefully acknowledged.

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Michener, C.D., and R.W. Brooks. 1984. Comparative
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Michener, C.D., and A. Fraser. 1978. A comparative an-
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Moure, J.S. 1944. Apoidea da cole^ao do Conde Amadeu
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. 1946. Notas sobre as Mamangabas. Boletim Agri-
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descritas por Perty em 1833. Boletim da Universidade
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. 1960b. Notes on the types of Neotropical bees

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Accepted 15 July 1985.

34 Contributions in Science, Number 369

Snelling and Brooks: The Tribe Ericrocini

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l'T 1 , -I ’ Vv ■ T

THE STATUS OF SMILODON IN NORTH AND SOUTH AMERICA

Annalisa Berta1 *

ABSTRACT. South American representatives of the sabercat genus
Smilodon (Felidae, Machairondontinae, Smilodontini) are referred
to a single species, Smilodon populator Lund, 1842, from Pleistocene
(?Uquian, Ensenadan-Lujanian) localities in Argentina, Bolivia,
Brazil, Ecuador, Peru, and Uruguay. North American Smilodon from
Pleistocene (late Irvingtonian-Rancholabrean) deposits are also as-
signed to -S’, populator. Diagnostic characters of A. populator include:
large size; skull with broad muzzle; upper canines strongly recurved;
upper canines and cheek teeth with finely serrated anterior and pos-
terior margins; P4 with very reduced protocone; mandible usually
with single, large mental foramen; mandibular flange greatly reduced.
Variation was noted in several characters; smaller size, triangular
occiput, and P3 were more common among middle Pleistocene (En-
senadan and Irvingtonian) samples of V. populator. During the late
Pleistocene, South American populations displayed a larger mean
size than their North American contemporaries.

Smilodon gracilis (Cope, 1880) is a more primitive related form
known from the late Pliocene (Blancan) through the middle Pleis-
tocene (Irvingtonian) of North America. The derived species S', pop-
ulator, apparently evolved in North America during the middle
Pleistocene (late Irvingtonian) and spread into South America. As
in North America, Smilodon became extinct in the southern con-
tinent at the end of the late Pleistocene (Rancholabrean and Luja-
nian, respectively).

INTRODUCTION

Although Smilodon has been known from the Pleistocene of
North and South America since the early part of the 19th
century, no previous studies have critically compared sam-
ples from the two continents. Such comparisons are essential
to an understanding of the phylogenetic interrelationships of
this sabercat and they also serve to strengthen existing bio-
stratigraphic correlations between the Americas during the
Pleistocene. The present contribution is an evaluation of the
status of Smilodon in North and South America with em-
phasis upon its South American record of occurrence.

ORGANIZATION AND METHODS

In addition to citations of the original publications of
F. Ameghino and P.W. Lund, cross-references to compila-
tions of their publications are given. The Obras Comp/etas
y Correspondencia de Florentino Ameghino edited by A.J.
Torcelli consists of 24 volumes published between 1913 and

Contributions in Science, Number 370, pp. 1-15
Natural History Museum of Los Angeles County, 1985

1926. The Memorias sobre a paleontologia Brasileira Revis-
tas e comentadas por Carlos de Paula Couto is a single vol-
ume of Lund’s work published in 1950. These compilations
include reproductions of most of the work of F. Ameghino
and P.W. Lund, and have had wider distribution than many
of the original publications.

The chronology and usage of South American late Ceno-
zoic Land Mammal Ages follows that proposed by Patterson
and Pascual (1972). From oldest to youngest, the Uquian,
Ensenadan, and Lujanian Land Mammal Ages are presently
included within the Pleistocene, with the Uquian straddling
the boundary between the Pliocene and Pleistocene. Corre-
lation of North and South American late Cenozoic Land
Mammal Ages follows Marshall et al. (1982) and suggests
the following correlations: Lujanian with late Ranchola-
brean, Ensenadan with early and middle Rancholabrean and
middle and late Irvingtonian, and Uquian with late Blancan
and early Irvingtonian.

Metrical data presented in this study should be interpreted
with caution. The scope of this analysis necessitated the use
of measurements published by others, although care was tak-
en to eliminate those that were not directly comparable with
the system of measuration utilized here and described by
Merriam and Stock (1932). Samples were often very small;
this was especially true for South American late Pleistocene
populations which unlike their North American contempo-
raries were not homogeneous with respect to locality and in
some cases possibly age. Statistics were computed through
the use of the BMDP (Dixon and Brown, 1979) programs.

The following institutions are referred to in the text: AMNH,
American Museum of Natural History, New York, New York;
BM(NH), British Museum (Natural History), London, En-
gland; FMNH, Field Museum of Natural History, Chicago,
Illinois; MACN, Museo Argentino de Ciencias Naturales
“Bernardino Rivadavia,” Buenos Aires, Argentina; MLP,
Museo de La Plata, La Plata, Argentina; MMP, Museo Mu-
nicipal de Ciencias Naturales y Tradicional de Mar del Plata,
Mar del Plata, Argentina; MNHN, Museum National d’His-
toire Naturelle, Paris, France; ROM, Royal Ontario Mu-

1 . Department of Biology, San Diego State University, San Diego,

California 92182-0063.

ISSN 0459-8113

seum, Toronto, Ontario, Canada; UZM L, Peter W. Lund
Collection, Universitets Zoologiske Museum, Copenhagen,
Denmark.

HISTORICAL REVIEW

The discovery of sabercats in South America was first an-
nounced by Lund (1842) and was based on material re-
covered from late Pleistocene-Holocene cave faunas of La-
goa Santa in eastern Brazil. Smilodon and a single species,
S', populator, were named from isolated teeth and associated
metapodials. Earlier, Lund (1839a) erroneously referred some
of this material to Hyaena neogaea. Since at the time of the
original designation Hyaena neogaea was insufficiently de-
fined and no type material was designated, this species is
regarded as a nomen nudum following Paula Couto (1955).

Subsequently, additional sabercats of Pleistocene age were
discovered in Argentina, Bolivia, Ecuador, and Peru. Most
of these specimens were attributed to Smilodon, although a
few were described as new genera and subgenera [i.e. Smi-
lodon ( Prosmilodon ) Rusconi, 1929; Smilodontidion Krag-
lievich, 1948],

Mendez-Alzola (1941) provided the most detailed descrip-
tion of South American sabercats and referred the Argentine
material to S. bonaeriensis. Paula Couto (1955) reviewed the
literature and, on the basis of Brazilian material, distin-
guished at the subspecific level the South American S. pop-
ulator populator from the North American S. populator cal-
ifornicus. Churcher (1967) examined some Argentine material
and added supplementary information about the South
American species which he referred to 5. neogaeus. Hoff-
stetter (1952) contributed to our knowledge of Smilodon from
Ecuador, while Churcher (written comm., 198 1) is describing
an excellent sample from the Talara tar seeps of Peru.

At present two species of Smilodon are recognized. The
small, slender S. gracilis is recorded from late Pliocene-mid-
dle Pleistocene (late Blancan-late Irvingtonian) localities in
North America (Berta, in press). The large, robust, and more
derived S. populator (synonyms include S. floridanus Leidy,
1889, S. californicus Bovard, 1907, and S. fatalis sensu
Slaughter, 1963) is recorded from middle-late Pleistocene
(late Irvingtonian-Rancholabrean) localities in North and
South America. Various authors (e.g. Webb, 1974b; Rob-
ertson, 1 976) have suggested interrelationships between North
and South American Smilodon, but until now such proposals
have been based on comparisons made from the literature
rather than firsthand study of collections.

GEOGRAPHIC AND STRATIGRAPHIC
DISTRIBUTION

Sabercats have been recorded from Argentina (Buenos Aires,
and Mar del Plata and environs), Bolivia (Tarija, Nuapua),
Brazil (Lagoa Santa Caves), Ecuador (La Carolina, Punin),
Peru (Talara), and Uruguay (Fig. 1). The oldest record, Smi-
lodontidion riggii (Kraglievich, 1 948), was reported from the
Chapadmalal Formation near Mar del Plata, Argentina, and
assigned a Chapadmalalan (late Pliocene) age. However, ac-
cording to G. Scaglia (pers. comm., 1981; Churcher, 1977:

255) this material was not collected in situ but rather from
landslide deposits at the base of a cliff that are more likely
Ensenadan in age.

The Uquian record of sabercats is based on material col-
lected from the Vorohue Formation at Mar del Plata and
Miramar, Argentina. According to J.L. Kraglievich’s (1952)
chronological subdivisions, this rock unit encompasses the
Vorohuean subage, an intermediate subdivision between the
Uquian and Ensenadan Land Mammal Ages, regarded by
Pascual et al. (1966) as indistinguishable from the Uquian.
Scaglia (pers. comm., 1981), however, questions the validity
of this record and interprets this material as having been
derived from Ensenadan-age deposits.

Discounting these questionable records, Smilodon is well
documented from deposits that range in age from Ensena-
dan-Lujanian. Argentine localities assigned an Ensenadan
age are listed by Mendez-Alzola (1941). Outside of Argen-
tina, sabercats have been recovered from middle and late
Pleistocene deposits of Tarija and Nuapua, Bolivia (Boule
and Thevenin, 1920; Hoffstetter, 1963, 1968). Known col-
lections of these faunas come from various stratigraphic levels
(MacFadden et al., 1983) and are Ensenadan and possibly
Lujanian in age. Lujanian age deposits that have yielded
abundant Smilodon include Buenos Aires, and Mar del Plata
and environs (Mendez-Alzola, 1941); Talara, Peru (Church-
er, written comm., 1981); La Carolina and Punin, Ecuador
(Hoffstetter, 1952); and various sites in Uruguay (Mones and
Francis, 1 973). Smilodon apparently became extinct in South
America during the late Pleistocene; the Lujanian-Holocene
Lagoa Santa Caves of Brazil represent its latest known oc-
currence.

SYSTEMATIC PALEONTOLOGY

Order Carnivora Bowdich, 1821

Suborder Feliformia Kretzoi, 1945

Family Felidae Gray, 1821

Subfamily Machairodontinae Gill, 1872

Tribe Smilodontini Kurten, 1963

Genus Smilodon Lund, 1 842

Hyaena Lund, 1839a:94 (not Hyaena Brisson, 1762).

Felis Lund, 1 839c:27 1 (not Felis Linnaeus, 1758).

Munifelis Muniz, 1845 (fide F. Ameghino, 1889:333).

Felis ( T rucifelis) Leidy, 1868:175.

Trucifelis Leidy, 1869:366.

Machaerodus Lydekker, 1884:33.

Drepanodon Leidy, 1889:14.

Machairodus Leidy, 1889:14.

Machaerodus Winge, 1895:3, 13 (not Machairodus Kaup,
1833).

Smilodontopsis Brown, 1908:188.

Prosmilodon Rusconi, 1929:5 (as a subgenus of Smilodon ).
Smilodontidion J.L. Kraglievich, 1948:6.

TYPE SPECIES. Smilodon populator Lund, 1842.

2 Contributions in Science, Number 370

Berta: Smilodon in the Americas

INCLUDED SPECIES. Type species and S. gracilis (Cope,
1880).

EMENDED DIAGNOSIS. Differs from all other ma-
chairodont genera but shares with Megantereon the following
derived characters: glenoid process enlarged; prominent post-
orbital process; well-developed supraoccipital crest; upper
canines elongate, compressed and recurved; upper and lower
incisors enlarged; P2/2 lost; long postcanine diastema; P3/3
reduced; P4 protocone reduced; lower canines reduced; and
limbs and feet short and stocky. More derived than Megan-
tereon in having incisors enlarged and procumbent; upper
canines longer and more robust; P4 ectoparastyle enlarged;
and mandibular flange reduced.

KNOWN DISTRIBUTION. ?Uquian. Ensenadan-Luja-
nian (middle-late Pleistocene) of South America and Irvmg-
tonian-Rancholabrean (middle-late Pleistocene) of North
America.

Smilodon populator Lund, 1842b

Figures 2-4

Felis cultridens Bravard, 1828:143, table III, figs. 10-13;

Blainville, 1864, II (Des Felis, Chap. 3:141).

Hyaena neogaea Lund, 1839a:94; 1 8 39b: 224. 232; 1840a:
265; 1840b: 3 12; 1842a: 12; Burmeister, 1864:123; 1879:
106; Paula C'outo, 1940:24; 1945:253.

Felis protopanther Lund, 1839c:207; 1840a:293, pi. XXVI,
figs. 10-11; Burmeister, 1879:106; H. Gervais and F.
Ameghino, 1880: (Obras 11:532).

Munifelis bonaerensis Muniz, 1845.

Machaerodus neogaeus{ Lund) Pictet, 1853:221; Burmeister,
1864:123; 1867:183; 1879:106; Lydekker, 1885:47; Winge,
1895:3, 13, 80, 82; Spillman, 1931:53; 1938:373.

Felis smilodon Desmarest, 1853:150, 152, 221, 223; 1860:
61; Blainville, 1855, pi. XX; Paula Couto, 1940:24; 1945:
253.

Smilodon blainvillii Desmarest, 1860:61.

Felis ( Trucifelis ) fatalis Leidy, 1868:175.

Trucifelis fatalis Leidy, 1869.

Machaerodus necator P. Gervais, 1878:1361 ( nomen nu-
dum).

Machaerodus cf. M. neogaeus Branca, 1 883: 1 37- 1 47, pi. XIX.
Machaerodus fatalis Lydekker, 1884:333.

Smilodon ensenadensis F. Ameghino, 1 888:6 (Obras V:473);

1898:195 (Obras XII:628); Paula Couto, 1945:253, 255.
Drepanodon or Machaerodus floridanus Leidy, 1889:14.
Machaerodus ensenadensis F. Ameghino, 1889:339 (Obras
XIL849, pi. IV, fig. 18); Bosca, 1923:182.

Smilodon floridanus Adams, 1896:433.

Smilodon crucians F. Ameghino, 1904:123 (Obras XV:201);

1 909: 122 (Obras XVII:610); l.L. Kraglievich, 1947:4, fig. 1.
Smilodon bonaerensis F. Ameghino, 1907:39, fig. 3 (Obras
XVII: 11, fig. 3); Rusconi, 1931:3, 4, 6, 8, figs. 1, 2, 4, 6,
7; Mendez- Alzola, 1941:10.

Smilodon californicus Bovard, 1907:155.

Smilodontopsis troglodytes Brown, 1908:188.
Smilodontopsis conardi Brown, 1908:190.

Figure 1. Distribution of Smilodon populator in South America.
Symbols: •, Lujanian; ■. Ensenadan; O, ?Uquian.

Smilodon nebraskensis Matthew, 1918:228.

Smilodon neogaeus ensenadensis Boule and Thevenin, 1920:
235, pi. XXVI, figs. 1-5.

Smilodon (Prosmilodori) ensenadensis Rusconi, 1929:5; 1936:
206.

Smilodon ( Trucifelis ) californicus Merriam and Stock, 1932:
16.

Smilodon ( Trucifelis ) fatalis Merriam and Stock, 1932:16.

Smilodon (Trucifelis) floridanus Merriam and Stock, 1932:
16.

Smilodon (Trucifelis) nebraskensis Merriam and Stock, 1932:
16.

Smilodon (Trucifelis) californicus brevipes Merriam and Stock,
1932:161.

Smilodon neogaeus Paula Couto, 1940:24-27; 1945:253, 255—
256, 261, note 38; 1950:58-59, note 18, 366, note 433,
538; 1953:96, 99; Churcher, 1967:245.

Smilodon necator Paula Couto, 1940:24; 1945:253.

Smilodon ( Prosmilodori ) ensenadensis feroxJ.L. Kraglievich,
1947:5.

Smilodon ( Prosmilodon ) ensenadensis minor J.L. Kraglie-
vich, 1948:11; Rusconi, 1929:5; 1931:14.

Contributions in Science, Number 370

Berta: Smilodon in the Americas 3

Figure 2. Smilodon populator, MACN 46, from near Lujan, Bue-
nos Aires Province, Argentina. Lateral (A) and ventral (B) views of
skull. Scale = 10 cm.

Smilodontidion riggii J.L. Kraglievich, 1948:6, figs. 3, 5-6,
8-9; Pascual et al., 1966:59.

Smilodon sp. Spillman, 1948:261; Hoffstetter, 1949:8; Hoff-
stetter, 1952:165, figs. 39, 40, pi. V, fig. 8, pi. VI, figs.
1-8.

Smilodon populator populator Paula Couto, 1955:1 1, fig. 10.
Smilodon trinitensis Slaughter, 1960:487.

Smilodon fatalis Slaughter, 1963:74.

SYNTYPES. UZM L specimens, right I2, I3, fragmentary
right C1, left metacarpals, II, IV, and right metacarpal V;
from Lagoa Santa Caves, Brazil.

Type of S. bonaerensis. MACN 46, a nearly complete skel-
eton, from Lujan, Buenos Aires Province, Argentina.

Type of S. crucians. MACN 1460, right ramus with I3-
M,; from Tarija, Bolivia.

Type of S. ( Prosmilodon ) ensenadensis ferox. MACN 1457,
anterior portion of right ramus with symphyseal region, C
root, and P3_4 alveoli; from Tarija, Bolivia.

Type of S. ( P .) ensenadensis minor. MACN 16693, occip-
ital portion of skull; MACN 16692, right cuboid; MACN
16689, right and left humeri; MACN 16695, proximal and

distal portions of left femur; MACN 1 6697, proximal portion
of right radius; MACN 16691, proximal portion of right tibia;
MACN 16690, left astragalus; right calcaneum; and MACN
16694, right metatarsal III. Probably all of the same indi-
vidual; from Punta Hermengo, Miramar Province, Argen-
tina.

Type of Smilodontidion riggii. MACN 6802, patella, prox-
imal phalanges III, IV, V, femur, tibia, astragalus, calcaneum,
metatarsal IV, fibula, and ischium; from Chapadmalal For-
mation, Buenos Aires Province, Argentina.

EMENDED DIAGNOSIS. Differs from Smilodon gra-
cilis in having large, robust skull with broad muzzle, upper
canines strongly recurved, upper canines and cheek teeth with
finely serrated anterior and posterior margins, P4 with very
reduced protocone, mandible usually with a single, large
mental foramen, and mandibular flange greatly reduced.

SOUTH AMERICAN DISTRIBUTION. Ensenadan-
Lujanian, Bolivia; Lujanian, Brazil; Lujanian, Ecuador;
?Uquian, Ensenadan-Lujanian, Argentina; Lujanian, Peru;
and Lujanian, Uruguay.

SOUTH AMERICAN MATERIAL STUDIED. See Ap-
pendix 1.

DESCRIPTION

Smilodon populator is best known in North America from
the late Pleistocene tar pits of Rancho La Brea, California
(Merriam and Stock, 1932). The South American record of
this sabercat is described herein using the Rancho La Brea
sample as the standard of comparison.

SKULL. Cranial measurements are listed in Appendix 2A.
South American individuals are characterized by develop-
ment of very large, robustly proportioned skulls. As seen in
dorsal view, the reduced nasals display deeply notched an-
terior ends and the muzzle is short and broad. The well
inflated postorbital processes are characterized by a marked
convexity of their posterior border. The zygomata are deep
and widely arched (Fig. 2A). The sagittal crest rises promi-
nently behind the slightly convex frontal region. In ventral
view (Fig. 2B), the auditory bullae are slightly inflated and
comparatively longer than in the Rancho La Brea sample.
The enlarged mastoid process is closely appressed to the
postglenoid process.

Some variation is apparent in the shape of the occiput (Fig.
3A, B). Most specimens examined show development of a
rounded, convex occiput with a strong vertical keel on the
basioccipital— the typical condition in S. populator as noted
by Merriam and Stock (1932:32). However, several speci-
mens including MACN 8682 display narrower, more trian-
gular occiputs with less distinct vertical keels. Slaughter (1963)
noted that triangular occiputs were characteristically devel-
oped among North American pre-Wisconsinan (late Irving-
tonian and early Rancholabrean) Smilodon which he referred
to S. fatalis. While it is generally true that most specimens
that display a triangular occiput are Ensenadan or Irving-
tonian in age, several Lujanian and Rancholabrean speci-
mens (e.g. MACN 8661, Fig. 3A) also exemplify this con-
dition. It is also possible that the shape of the occiput may
represent sexual dimorphism or age variation as it was ob-

4 Contributions in Science, Number 370

Berta: Smilodon in the Americas

A

B

Figure 3. Smilodon populator, occipital views of skull showing
extremes in variation. A, MACN 8661 from Olivos, Buenos Aires
Province, Argentina; B, MACN 8682 from P. Piedras, Buenos Aires
Province, Argentina. Scale = 10 cm.

Figure 4. Smilodon populator, MACN 46, from near Lujan, Bue-
nos Aires Province, Argentina. Lateral (A) and occlusal (B) views of
mandible. Scale = 5 cm.

served that those specimens that exhibit a narrower, more
triangular occiput usually possess well-developed sagittal
crests and often typify subadult individuals.

UPPER DENTITION. Appendix 2B lists upper tooth di-
mensions. Upper incisors are large, sharp and procumbent.
I ' is comprised of lingual and labial cusps positioned on either
side of the V-shaped notch encircling the principal cusp. I2
is similar in morphology, with the lingual ridge extending
further anteriorly. The caniniform I3 consists of a principal
cusp and small basal cusp positioned on a circular serrate
ridge on the lingual tooth margin. The strongly developed
posterolateral ridge is serrated in unworn specimens.

The long, slender canine is strongly recurved and hne enamel
serrations are developed on the anterior and posterior mar-
gins. Although few sabers are preserved in entirety, the canine
is longer than in Rancho La Brea specimens (Fig. 2). Paula
Couto (1955, table 1) lists canine length for a specimen from
Brazil as approximately 277.5 mm compared with a range

Contributions in Science, Number 370

Berta: Smilodon in the Americas 5

of measurement for this tooth among the Rancho La Brea
sample of 200-262 mm (Merriam and Stock, 1932:48).

P3 is relatively small and variable in morphology. It is
double-rooted and consists of a principal cusp flanked by
anterior and posterior cusps. Occasionally, a second posterior
cusp develops, as in ROM 2116 and MLP 10-2. A strong
anterolingual cingulum is usually present. In MACN 46,
however, the anterior and posterior cusps are small, and the
anterolingual cingulum is absent. The orientation of P3 varies
from a linear to oblique position behind the canine.

The small ectoparastyle and parastyle of the upper car-
nassial form the anterior blade of the tooth. The metacone
is always anteroposteriorly longer than the paracone. The
protocone is usually very reduced or absent although a sep-
arate protoconal root is usually present. The occasional pres-
ence of the protocone was noted in Rancho La Brea Smilodon
by Merriam and Stock ( 1 932:48) and confirmed in this study.
Unworn specimens characteristically show development of
serrations along the cutting edge. M1 is a very reduced, dou-
ble-rooted tooth with a transversely elongated crown com-
prised of a principal cusp, the parametacone, and a small
protocone. The tooth is oriented at nearly right angles to the
anteroposterior axis of P4.

MANDIBLE. The horizontal ramus is anteroposteriorly
short and robust (Fig. 4A, B, Appendix 2A). The anterior
face of the massive symphysis is concave and pitted by fo-
ramina. The flange is very reduced ventrally in comparison
with Smilodon gracilis, although its anterior margin is usually
thick and laterally flared. The diastema between the C and
P4 is comparatively shorter among most of the South Amer-
ican sample although its length is variable, a feature appar-
ently related to age (Merriam and Stock, 1932, pi. 4). Usually
a single, large mental foramen is positioned below the dia-
stema near the ventral border of the ramus although double
foramina occasionally occur in the Californian and Talaran
populations (Churcher, 1984; this study). The coronoid pro-
cess is greatly reduced. Among several South American spec-
imens (e.g. MACN 46 and MACN 18057) the prominent
angular process swings laterally farther away from the deeply
excavated masseteric fossa than in most Rancholabrean spec-
imens (Fig. 4).

LOWER DENTITION. Appendix 2B lists lower tooth
dimensions. Lower incisors are broad, recurved, and set in
a transverse line. The small I, bears a prominent V-shaped
notch encircling the principal cusp and a lingual ridge which
terminates in a basal cusp. I2 is similar in morphology, with
serrations present on the lingual ridge. The caniniform I3
exhibits a prominent serrate lingual ridge. The lower canine
has in addition to the lingual ridge a prominent posterior
ridge, slightly labial to the midline of the tooth; both ridges
are serrated.

P3, commonly present in Ensenadan and Irvingtonian
specimens, is extremely variable in morphology (CV for
length = 23.4). In MACN 6270, P3 is single-rooted and com-
prised of a single principal cusp followed by a small posterior
cusp. In another specimen, ROM 5100, the alveolus for this
tooth indicates that it was double-rooted. Loss of P3 is more
common among Lujanian and Rancholabrean specimens. As

noted by Merriam and Stock (1932:5 1), this tooth is present
in only 6% of the La Brea sample (N = 678).

P4 is double-rooted and comprised of a principal cusp
flanked by a relatively small anterior cusp and two progres-
sively smaller posterior cusps. The second posterior cusp is
occasionally positioned on a strong lingual cingulum. The
crown of this tooth and M, are inclined posteriorly.

M, usually possesses a small anterior cingulum in front of
the paraconid. Presence of this cusp among North American
specimens was previously noted (Bovard, 1907). The para-
conid blade is noticeably shorter anteroposteriorly than the
protoconid and both are serrated. Most specimens show slight
development of a basal posterior cingulum behind the pro-
toconid.

DISCUSSION

The known South American samples of sabercats are all
referred to Smilodon populator. As previously mentioned,
Smilodon was established by Lund (1842b) who described a
single species, A. populator, from the Lagoa Santa Caves,
Brazil. Muniz (1845) described a sabercat skeleton collected
from near Lujan, Argentina, under the name Munifelis bo-
naerensis. However, his generic name is invalid since it was
published in a newspaper (La Gaceta Mercantil), which does
not satisfy the publication requirement established by the
International Code of Zoological Nomenclature. Later au-
thors (Ameghino, 1907; Rusconi, 1931; Mendez- Alzola, 1941)
incorrectly referred to sabercats under the name Smilodon
bonaerensis. Burmeister (1867) applied the name Machae-
rodus bonaerensis to the Argentine material, although he later
acknowledged that S. bonaerensis was probably conspecific
with S. neogaeus (Lund, 1839a).

Ameghino (1888:6) described Smilodon ensenadensis
(=Machaerodus ensenadensis) based on an upper canine re-
covered from Ensenadan deposits in Ensenada, Argentina.
Rusconi (1929) listed small sabercats from the Ensenadan of
Argentina under the subgenus Smilodon ( Prosmilodon ).
L. Kraglievich (1934) followed Rusconi’s usage of Prosmi-
lodon. In 1947 J.L. Kraglievich formally diagnosed Smilodon
( Prosmilodon ) on the basis of its small size and presence of
P3. The variable presence of this tooth among smilodontines
has been previously discussed. Presence or absence of this
tooth generally can be used to distinguish Ensenadan or Ir-
vingtonian 5. populator from Lujanian or Rancholabrean
forms of this species.

A right ramus (MACN 1457) collected from Tarija, Bo-
livia, and originally described and figured by Ameghino ( 1 902)
as Machaerodus ensenadensis, was referred by Kraglievich
(1947) to Smilodon (Prosmilodon) ensenadensis ferox. This
subspecies was distinguished from S. bonaerensis on the basis
of its smaller size, lack of serrations on the upper canine,
and presence of a double-rooted P3. Examination of the type
and referred material indicates that the upper canine is heavi-
ly worn along both the anterior and posterior margins, oblit-
erating serrations had they been present. A second subspe-
cies, Smilodon ( Prosmilodon ) ensenadensis minor, was
distinguished by J.L. Kraglievich (1948) on the basis of its

6 Contributions in Science, Number 370

Berta: Smilodon in the Americas

Table 1A. ANCOVA for length of lower canine of Smilodon pop-
ulator adjusted for length and width of P4 as a test for significant
differences based on locality and age. * = P < 0.05, ** = P < 0.01.
Abbreviations = AP = anteroposterior length, T = transverse width.

Souce of
variation

Sum of
squares

Degrees

of

free-

dom

Mean

square

F-value

Regression

coefficient

Locality

0.06274

1

0.06274

0.04

Age

37.41456

1

37.41456

23.32**

Locality/age

29.88608

1

29.88608

18.63**

AP, P4

6.25910

I

6.25910

3.90*

0.23330

T, P4

0.00007

1

0.00007

0.00

-0.00183

AP & T, P4

12.34655

2

6.17328

3.85*

Error

75.41 120

47

1.60449

smaller size and more robust postcranium. Examination of
the type indicates that it falls within the limits of variation
shown by Smilodon. The subgenus Smilodon ( Prosmilodon )
is here referred to Smilodon.

Smilodon crucians was recognized by Ameghino ( 1 904) on
the basis of its small size, shallow ramus, and long postcanine
diastema. These characters are attributable to ontogenetic
variation (see Merriam and Stock, 1932, pi. 4, fig. 10) and
indicate that S. crucians represents a juvenile individual. As
expected, the high coefficient of variation value for postca-
nine diastema length among both North and South American
sabercats (see Appendix 2A) further weakens its use as a
taxonomic character. Examination of the holotype of S. cru-
cians (MACN 1460) revealed the presence of serrations on
the lower canine and cheek teeth, which supports synonymy
of this species with S. populator.

A new genus of sabercat, Smilodontidion, was proposed
by J.L. Kraglievich (1948) on the basis of limb elements
collected from near Chapadmalal, Argentina. This genus was
distinguished by its smaller size and slightly different mor-
phology, regarded here as the result of both individual and
age variation. The incomplete fusion of epiphyses on the
femur and tibia indicates that this specimen represents a
young adult. I agree with Churcher (1967) that Smilodonti-
dion should be synonymized with Smilodon.

The results of systematic reevaluation of North American
Smilodon are summarized in the following discussion (see
also Berta, in press). The oldest name for a North American
Smilodon is Felis ( Trucifelis ) fatalis proposed by Leidy ( 1868).
The type, a fragmentary maxilla containing P4 from Hardin
County, Texas, redescribed and illustrated by Leidy (1869)
is judged too incomplete to provide specific diagnosis. Later,
Leidy (1889) described a fragmentary skull lacking teeth as
a new species of the European genus Machairodus, M. flor-
idanus. This species has been referred to Smilodon by all
later workers. Bovard (1907) proposed the species of Smi-
lodon californicus separating it from S. floridanus by the

Table 1 B. Adjusted group means for length of lower canine of Smi-
lodon populator. Abbreviations: N = sample size, as in Table 1A.

North America

South America

Age

Ranchoiabrean

Irvingtonian

Lujanian

Ensenadan

AP, C

14.81744

13.80000

17.81667

11.43333

N

43

4

3

3

latter’s shorter muzzle, more posterior position of the pos-
terior nares and possibly by ridges on the palate. The sample
from Rancho La Brea consisting of thousands of individuals
was more fully described by Merriam and Stock (1932) who
proposed certain differences in the mastoid region to distin-
guish the two species. Kurten’s (1965) study of Florida sa-
bercats led him to synonymize S. floridanus with S. fatalis
and maintain S. californicus as distinct. Webb ( 1974b) in his
evaluation of additional Florida material strengthened the
case for synonymy of S. californicus with S', floridanus using
broad overlap of cranial, mandibular, and tooth measure-
ments.

More recently, Kurten and Anderson (1980) recognized a
single derived species, Smilodon fatalis and considered “the
numerous North American taxa of sabertooths (with the ex-
ception of S. gracilis) to have, at most, subspecific standing.”
Churcher (1984) followed Slaughter (1963) in assigning to
Smilodon fatalis the following species: Smi/odontopsis co-
nardiBrown, 1 908; Smilodontopsis troglodytes Brown, 1908;
Smilodon nebraskensis Matthew, 1918; and Smilodon trin-
itensis Slaughter ( 1 960).

The genus Smilodontopsis proposed by Brown (1908) was
founded on specimens from the Conard Fissure, Arkansas,
and distinguished from Smilodon by the possession of an
entepicondylar foramen on the humerus. Two species, S.
troglodytes and S', conardi were described on differences in
the morphology of the upper camassial. Smilodon nebras-
kensis was based on a lower jaw from Nebraska and distin-
guished from S. californicus by its “decidedly shorter post-
canine diastema” (Matthew, 1918), a character of suspect
taxonomic value. Smilodon trinitensis was founded by
Slaughter (1960) on a lower jaw from the Trinity River, at
Dallas, Texas. It was distinguished from previously described
species by its large cheek teeth, relative to mandibular mea-
surements and the absence of P3. As noted previously, P3 is
more commonly present in middle Pleistocene samples and
is extremely variable in morphology.

The taxonomic confusion surrounding North American
Smilodon brought out in the preceding discussion suggested
as the next step, critical evaluation of the status of S. flori-
danus and S. fatalis (sensu Slaughter, 1963). As shown in
this study, all of the characters diagnostic of North American
S. floridanus also typify South American S. populator. Among
the most useful of these characters are: large, robust skull
with broad muzzle, upper canines strongly recurved, upper
canines and cheek teeth with finely serrated margins, P4 with

Contributions in Science, Number 370

Berta: Smilodon in the Americas 7

.08 .06 .04 .02 0 .02 .04 .06 .08

Figure 5. Ratio diagram of cranial, mandibular, tooth, and limb measurements (see Appendix 2) of South American Smilodon populator
(solid line, Lujanian sample; dashed line, Ensenadan sample) compared with North American Smilodon populator (Rancho La Brea sample;
Merriam and Stock, 1932) as the standard. A log difference scale is provided and horizontal bars show the observed range of variation of the
standard.

very reduced protocone, mandible usually with a single, large
mental foramen, and mandibular flange greatly reduced.

A ratio diagram (Fig. 5 see Appendix 2 for sample statistics;
for the method see Simpson, 1941) illustrates the differences
between mean measurements of South American S. popu-
lator and the Rancho La Brea sample, the standard of com-
parison. Two sets of data are compared: (1) skull and man-
dible dimensions and (2) forelimb and hindlimb lengths from
both North and South American middle and late Pleistocene
samples. South American specimens of S. populator from
late Pleistocene (Lujanian) localities are on the average larger
for nearly every dimension studied than North American
Rancho La Brea individuals, an observation noted previously
by Paula Couto (1955) and Kurten (1965). However, nearly
all of the South American specimens fall within the observed
range of variation for the Rancho La Brea sample.

Although sample sizes are small, mandibular dimensions
of middle Pleistocene (Ensenadan) S. populator average
smaller than either Lujanian or Rancholabrean specimens.
Mean values of lower tooth dimensions for South American
S. populator from Ensenadan localities approximate those of
North American Irvingtonian individuals with several ex-
ceptions. Among Ensenadan individuals P3 averages larger
whereas P4 and M, average smaller than corresponding teeth
in Irvingtonian specimens (Appendix 2B).

While most limb elements are short and stocky in smilo-
dontines, this is not true for the femur of S. populator which
is proportionately longer and not nearly as robust as in most
large felids (Gonyea, 1976, table 4, fig. 6). Analysis of limb
lengths (Appendix 2C, Fig. 5) suggest that the principal fore-
and hindlimb elements (humerus, femur, and tibia) average
longer among Lujanian S. populator than the same elements
among Rancholabrean individuals. Proportions of the hind-
limb of Ensenadan S. populator are much the same as the
Rancholabrean population except for their smaller size.

Slaughter (1963) used five characters to diagnose 5. fatalis.
Each of these characters is listed below and then reconsidered
in light of reexamination of both North and South American
specimens.

(1) Total skull length less than any specimen of S. calif or -
nicus (=S. floridanus). Following Merriam and Stock (1932),
Slaughter (1963, table 3) lists the smallest skull from Rancho
La Brea as 256.4 mm. Although the present study substan-
tiates this claim, six skulls ranging between 259.5 and 273.0
mm increase the previously known North American sample
of “small” S. populator individuals.

(2) Occiput narrow and triangular, not rounded above. As
previously discussed, occiput shape is variable. North and
South American populations of middle and late Pleistocene
S. populator contain specimens which exhibit narrow, tri-

8 Contributions in Science, Number 370

Berta: Smilodon in the Americas

angular occiputs although they are more common among
middle Pleistocene individuals.

(3) Basioccipital lacking well-developed keel. This was the
most variable character observed. Although it is often as-
sociated with a more triangular occiput, just as many indi-
viduals with a rounded occiput display an indistinct, flattened
keel.

(4) Inferior canine much reduced with anteroposterior di-
ameter less than 54% of the same measurement of P4.

(5) P4 thin, having a transverse diameter less than 43% of
the anteroposterior diameter. In his figures 1 and 2 Slaughter
showed a difference between North American Smilodon cal-
ifornicus material of late Pleistocene (Wisconsin) age from
California and Florida and a smaller group of all middle
Pleistocene (pre-Wisconsin) material which he referred to
Smilodon fatalis. Figure 6 presented here repeats Slaughter’s
bivariate plots with the addition of South American Smi-
lodon populator. When these data are added the ranges of
the various measurements overlap and strengthen the case
for synonymy of North American Smilodon species with
South American Smilodon populator.

Slaughter’s dental characters (length of lower canine ad-
justed for length and width of P4) were further evaluated in
an analysis of covariance (ANCOVA) for Smilodon populator
collected from (1) different localities (North vs. South Amer-
ica) and (2) different ages (middle vs. late Pleistocene) (Table
1A and B). Mean length of canine for S. populator differed
only in age within the South American sample which led to
the significant interaction term (Table IB). However, for the
characters evaluated sample sizes for middle Pleistocene
Smilodon are very small (N = 4, North America; N = 3, South
America). When age is eliminated as a factor and P4 length
is adjusted for width, using a pooled late Pleistocene sample
there is no significant difference in adjusted means between
North and South American S. populator (adjusted group
means for length of P4; North American samples = 39.74772,
South American samples = 4 1 .48333). The results of analysis
of covariance using Slaughter’s dental characters do not sup-
port recognition of more than one Smilodon species; both
North and South American samples are referred to S. pop-
ulator.

The practice of recognizing a species solely on the basis of
size is questioned here for several reasons. Earlier work
(Slaughter, 1963; Webb, 1974b) has shown that size overlap
in a number of cranial and tooth measurements for Smilodon
is too great to permit species discrimination. Size variation
has been previously identified among wide-ranging large car-
nivores; the spotted hyena (Crocuta crocuta ) (Kurten, 1957)
and the puma (Felis concolor) (Kurten, 1973). Strong sexual
dimorphism in the puma was demonstrated for all skull and
tooth measurements studied (condylobasal length, zygomatic
width, C length, P4 length, and M, length). The results of
comparison of these variances in a standardized local pop-
ulation of North American Felis concolor statistically ad-
justed for 35°N latitude (Kurten, 1973; table 2) and in the
combined North and South American Smilodon populator
are given in Table 2. Statistically significant differences were

18-

17-

16-

15-

0

1 14-

Q 13-

Q-

<

12

11-

d

101

22 23 24 25 26 27

AP Diameter P4

28 29 30

Figures 6A, 6B. Fig. 6A. Bivariate plot of canine length versus P4
length for Smilodon populator and Smilodon fatalis (sensu Slaughter,
1963). Plot gives comparison of S. populator from North America
(data from Merriam and Stock, 1932 and this study; •, Ranchola-
brean) and South America (data from this study; O, Lujanian, d
Ensenadan, and Smilodon fatalis from North America (data from
Slaughter, 1963; ■, Rancholabrean, d, Irvingtonian). Fig. 6B. Bi-
variate plot of P4 width versus length for Smilodon populator and
Smilodon fatalis (sensu Slaughter, 1963). Comparison and symbols
as above.

observed between these variances for only two dimensions,
condylobasal length and zygomatic width. If one assumes a
similar pattern of variation among other large cats then great-
er importance can be attached to the fact that all other di-
mensions (C, P4, and M , lengths) do not show any statistically
significant differences between samples. The data indicate
that for these tooth dimensions, variation in S. populator can
be accounted for simply as the result of sexual dimorphism.
Kurten was able to distinguish sexual variation from indi-
vidual and latitudinal variation in the puma. In the present
study comparisons were made on the basis of total variability
within S. populator since I could not partition the variances
associated with either sex or latitude. Based on Kurten’s data.

Contributions in Science, Number 370

Berta: Smilodon in the Americas 9

Table 2. Variation in Felis concolor' and Smilodon populator from
North and South America. Abbreviations: N = sample size; s2 =
variance; F = F ratio; P = probability; 1 = data from Kurten, 1973,
table 2.

Character

N

Taxon

s2

F

P

Condylobasal

116

F. concolor

163.890

1.928

-0.02

length

28

S. populator

316.008

Zygomatic width

118

F. concolor

1 15.313

8.170

-0.001

30

S. populator

14.1 13

C length

144

F. concolor

1.507

1.435

-0.2

37

S. populator

2.164

P4 length

141

F. concolor

0.649

1.200

-0.5

31

S. populator

0.79

M, length

143

F. concolor

0.268

1.06

-0.5

42

S. populator

0.252

undoubtedly, both individual and latitudinal variation are
important aspects to the observed variation in Smilodon.
However, the fact that the comparable variation in tooth
dimensions in North and South American Smilodon can be
accounted for solely by sexual dimorphism without consid-
eration of these other parameters, further strengthens the
argument for recognition of Smilodon populator as a single
wide-ranging species.

PHYLOGENETIC RELATIONSHIPS AND
ZOOGEOGRAPHIC PATTERNS

Judging from the oldest stratigraphic record, sabercats of the
genus Smilodon originated in North America during the late
Cenozoic. S. gracilis (Cope, 1880), the sister taxon of the
more derived S', populator, is well represented from the late
Blancan through Irvingtonian of Florida. This species is also
known from the type locality at Port Kennedy Cave in Penn-
sylvania and with less certainty from the Palm Springs For-
mation, Vallecito Creek, California (Kurten and Anderson,
1980; Berta, in press). The following characters distinguish
S. gracilis from S. populator. canines only slightly recurved;
canines and cheek teeth with very finely serrated margins; P4
with well-developed protocone; mandible with two mental
foramina and an enlarged flange (Berta, in press).

The earliest record of S. populator is from the Irvingtonian
of Arkansas, California, and Nebraska. During the late Pleis-
tocene S. populator ex tended its range throughout the United
States, Central America, and into South America. In South
America, this species has been recorded from Argentina, Bo-
livia, Brazil, Peru, and Uruguay. The extension of 5. popu-
lator into South America is not so surprising when one con-
siders the equally wide range of such present-day species as
the lion, Felis leo and the puma, Felis concolor.

The end of the Pleistocene marked the extinction of Smi-
lodon in both North and South America. As was true for
large canids the extinction of sabercats can be related to
extinction of their large, thick-skinned herbivorous prey which

included ground sloths, glyptodonts, proboscideans, horses,
notoungulates, and litoptems. While supporting this inter-
pretation of Smilodon prey, Akersten (1985) added that the
“canine shear-bite” of this species would have been just as
effective on smaller thin-skinned prey. However, it should
be stressed, as elaborated by Rosenzweig (1966), that larger
carnivores generally take larger prey. It seems likely that the
extinction of sabercats can be related in part to their spe-
cialized hunting and feeding strategy and resultant social be-
havior which was dependent on a large herbivore megafauna,
so reduced in number and diversity at the present time.

ACKNOWLEDGMENTS

Much of this research was undertaken at the Vertebrate Pa-
leontology Section of the Museo Argentino de Ciencias Na-
turales “Bernardino Rivadavia,” Buenos Aires (MACN),
where Jose Bonaparte (Chief of Section of Fossil Vertebrates)
made collections, working space, and library facilities avail-
able. Rosendo Pascual and Guiomar Vucetich, Facultad de
Ciencias Naturales y Museo de la Universidad Nacional de
La Plata (MLP), and Galileo J. Scaglia, Director, Museo
Municipal de Ciencias Naturales, Mar del Plata (MMP), gave
permission to study collections under their care and facili-
tated research in many ways. David Wright measured spec-
imens in the University of Nebraska collections. D.A. Farris
assisted with statistical manipulation of the data. W. Aker-
sten, H. Galiano, E. Manning, B.J. MacFadden, L. Marcus,
C.A. Repenning, R.G. Wolff, and S.D. Webb critically re-
viewed this manuscript.

Financial support from Sigma Xi, the Hays Fund of the
American Philosophical Society, Thomas J. Dee Research
Fellowship (Field Museum of Natural History, Chicago), the
Department of Natural Sciences, Florida State Museum,
University of Florida is gratefully acknowledged. This re-
search was initiated during tenure of a Postdoctoral Fellow-
ship at the Department of Natural Sciences, Florida State
Museum, University of Florida. This report is University of
Florida Contribution to Vertebrate Paleontology number 214.

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Accepted 15 July 1985.

APPENDIX 1.

SOUTH AMERICAN MATERIAL STUDIED
ARGENTINA

MACN Specimens: 46, nearly complete skeleton (type of S. bona-
erensis)-, 997, incomplete skull; 1107 humerus; 2354, right ramus
with I|_3, C, left tibia; 2838, humerus; 5010, right ramus; 5109, right
femur; 5273, skull, right ramus, proximal femur, tibia, ulna; 5822,
fragmentary skull (juvenile); 6205, left ramus with C root and P4-
M,; 6210, right ramus with P3-M,; 6234, tibia; 8661, posterior
portion of skull; 8682, posterior portion of skull; 8707, right femur;
9446, distal right humerus; 9650, right humerus; 10037, proximal
femur; 10485, tibia; 10502, proximal humerus; 10523, radius; 10861,
ulna; 1 1554, right ramus with I3, C alveolus, P4-M,; 12011, rostral
fragment; 12393, fragmentary lower I; 12445,C, ; 1 2474, C fragment;
12532, ramus with M,_2; 12846, skull fragment; 13606, C1; 13922,
occipital fragment; 14165, left maxillary fragment with P3-4; 13206,
M,; 16693, occipital portion of skull; 16692, ulna; 16689, right and
lefthumeri; 16695, proximal and distal portions of left femur; 16697,
proximal right radius; 16691, proximal right tibia; 16690, left as-
tragalus; 16694, right metatarsal III; [12846-16694 are probably all
from the same individual, the type of A. (P.) ensenadensis minor;]
17448, fragmentary C\ incomplete skull; 18026, right maxillary
fragment with P4; 18051, symphyseal portion of mandible; 18057,
skull and mandibles.

MLP Specimens: 10-1, skull and mandibles; 10-2, incomplete
skull and mandibles; 10-10, right ramus; 10-13, left humerus; 10-
14, left humerus; 10-20, left ulna; 10-22, right radius; 10-23, right
radius; 10-33, left tibia; 10-35, left tibia; 10-37, incomplete right
scapula; 10-44, pelvis, sacrum, and lumbar vertebrae; 10-47, axis;
10-48, thoracic and lumbar vertebrae, sacrum, pelvis, right femur.

MMP Specimens: 5M, left femur; 2 1 -S, posterior portion of skull;

Contributions in Science, Number 370

Berta: Smilodon in the Americas 13

34-S, left hindlimb (distal tibia and fibula, astragalus, calcaneum,
mesocuneiform, ectocuneiform, cuboid, navicular, metatarsal II-V,
proximal phalanges and digits I-V, medial phalanges and digits II-V ;
ungual phalanges I-IV, sesamoids); 227-S, proximal femur; 465-S,
left femur; 495-S (a-f), left and right tibiae, distal epiphysis of femur,
astragalus, cervical vertebrae V-VI; 652-S, humerus; 765-M (a-g),
phalanges; 768-M, M,, left P4; 796-M (a-w), portion of skull, ver-
tebrae (atlas, lumbar, sacrum), pelvis, hindlimb (distal femur, fibula,
tibia, left and right astragalus, right and left calcaneum, ectocunei-
form, cuboid, navicular, right and left metatarsals II, III, IV, V);
1 3 1 1 -M, incomplete skull with right maxillary, parietal and occipital
regions partly restored.

AMNH Specimens; 11101, skull and mandible with nearly com-
plete skeleton; 11103, vertebrae and skeletal fragments; 1 1 104, pel-
vis; 1 1 105, left femur; 1 1 106, right calcaneum.

FMNH Specimens: P 14294, complete mandible and partial skel-
eton including: atlas, cervical and thoracic vertebrae, ribs, sternum-
brae, humeri, left scapula, ulna, radius, scapholunar, unciform, mag-
num, metacarpal I, proximal, medial and ungual phalanges, pelvis,
femora, tibiae, astragali, left metatarsal II, right metatarsals III, IV
and V, and patella; P 14271, skull and mandible (partially restored)
and partial skeleton including: atlas, cervical, thoracic, and lumbar
vertebrae, sacrum, ribs, left humerus, left and right metacarpal II,
right metacarpals II and V, left scapholunar, proximal phalanges,
pelvis, right fibula, patella, left astragalus and calcaneum; P 14279,
left maxillary fragment with P3^.

BOLIVIA

MACN Specimens: 1455, left premaxillary and maxillary fragment
with I2-3, C; 1457, anterior portion of right ramus with C, P3 alveolus,
P4 (type of S. (P.) ensenadensis ferox)\ 1459, distal left humerus;
1460, right ramus with I3-M,, right and left humerus, left tibia,
manus (type of S. crucians).

BRAZIL

UZM L Specimens: right I2, 13, fragmentary right C\ left metacarpal
II and IV, right metacarpal V (type of S. neogaeus)\ left distal meta-

carpal II, phalanges (hindfoot), C1, right ramus with incisor alveoli,
C root, P4-M, (text and figures in Paula Couto, 1955); Winge (1895:
13) lists additional cranial and postcranial material housed in the
UZM collections.

BM(NH) Specimens: 18972, left P3~4 associated with skull figured
by Blainville (1855), left femur (diaphysis), right humerus, right ulna,
right distal tibia, right astragalus, phalanx, cervical vertebrae, and
right calcaneum, all probably from the same individual.

MNHN, right side of skull with I'-3, P4-M‘, left C, and fragmen-
tary ramus with complete dentition (text and figures in Blainville,
1855).

ECUADOR

EPN Specimens: V. 1507, nearly complete skull; V. 1903, cervical
vertebrae; V.2990, anterior portion of right scapula; V. 1902, prox-
imal right radius; V. 1 899, left magnum; V. 1 89 1 , right patella; V. 1 894,
right tibia; V. 1895, right tibia; V. 1896, proximal right tibia; V.1212,
right cuboid; V. 1893, left metatarsal III; V.l 182, right C1; V.l 185,
fragment of left maxillary with P3-4; V. 1 1 86, maxillary fragment with
right I1-2, left I1-3; V.l 187, fragment of right maxillary with P4; V.l 193,
right I3; V.l 197, fragmentary right I3; V.l 195, left I2; V.l 183, man-
dible with left I2_3, P4-M, and right I2_3, C, P4-M,; V.l 184, right
ramus with P3 alveolus; V.l 191, left dP4 in fragmentary mandible;
V.718, fragment of scapula; V.l 188, right humerus; V.l 189, left
scapholunar; V. 1 1 90, left metacarpal IV and V, right metacarpal V;
V.l 206, left metacarpal III; V.l 203-04, phalanx I, digit I (forefoot);
V.l 292, patella; V.l 180, left “perone”; V.l 198-1202, phalanges (text
and figures in Hoffstetter, 1952).

PERU

The extensive Talaran collection of S. populator is currently being
described by C.S. Churcher (written comm., 1981).

14 Contributions in Science, Number 370

Berta: Smilodon in the Americas

Appendix 2A-C. Measurements and statistics for Smilodon populator from North and South America. Abbreviations: a, approximate mea-
surement; AP, greatest anteroposterior dimension; CV, coefficient of variation; N, sample size; OR, observed range of variation; SE, standard
error; T, greatest transverse dimension; x, mean; 1, measurements following Mendez-Alzola, 1941; Hoffstetter, 1952; Churcher, 1967 and this
report; 2, measurements following Merriam and Stock, 1932; 3, measurements following Slaughter, 1963 and this report.

2A. Cranial ar

id mandibular measurements

South America1

North America2

Ensenadan

Lujanian

Rancholabrean

N

x ± SE

OR

CV

N x ± SE

OR CV

N

x ± SE

OR

CV

1. Length, premaxillary-condyle —

— -

-

3 358.6 ±

3.18

355.0-365.0 1.54

25 308.6 ± 0.958

271.4-344.1

7.77

2. Length, premaxillary-inion

— -

-

4 352.3 ±

10.46

335.0-353.0 5.93

25 336.0 ±1.000

292.7-377.9

7.45

3. Length, C1

-P4 1

1 10 -

-

3 121.2 ±

0.186

121.1-

-121.6 0.264 24 112.1 ±0.330

97.0-126.3

7.08

4. Greatest width, zygomatic arches —

- -

-

6 211.8 ±

5.49

191.0-225.0 0.634 24 208.8 ± 0.626

182.9-240.1

7.20

5. Palatal width at P4

— -

—

4 128.6 ±

5.06

117.0-140.0 7.86

24 127.5 ±0.298

i 14.7-135.4

5.62

6. A-P diameter, auditory bulla —

- -

-

4 60.3 ±

2.89

53.7-67.8 9.59

24

54.2 ± 0.224

46.7-63.9

9.93

7. Greatest width, mastoid processes 3

136.9 ± 3.89

132.0-144.6

4.92

9 138.9 ±

3.71

1 14.4— 147 6 8.01

24 134.6 ±0.431

122.0-154.3

7.71

8. Greatest width, condyles 3

70.2 ± 3.02

66.1-76.0

7.33

10 71.7 ±

1.33

66. 1-

-78.1 5.86

25

65.0 ± 0.154

57.6-72.3

5.94

9. Length, symphysis-condyle 2

194.5 -

183.0-206.0

-

9 224.3 ±

5.00

198.0-242.8 6.63

25 209.5 ± 0.632

178.3-230.0

7.55

10. Length, diastema (C,-P3) 3

54.7 ± 3.78

50.1-62.0

1 1.65

3 68.1 ±

39.32

64.7-

-70.0 4.37

25

61.7 ± 0.314

46.3-71.8

12.72

1 1. Depth, ramus below M, 3

36.8 ± 2.31

32.7-40.7

10.88

8 43.0 ±

15.21

40.0-49.0 7.81

25

40.3 ± 0.1 10

36.0-45.6

6.84

12. Mandibular toothrow length 2

50.5 -

48.0-53.0

-

3 52.7 ±

2.05

48.0-57.0 6.59

25

54.1 ± 0.123

48.3-60.9

5.72

13. Height, angle-coronoid 1

48.5 -

-

6 71.0 ±

3.60

61.1-

-84.2 12.43

25

68.9 ± 0.207

60.3-76.3

7.53

2B. Upper and lower tooth measurements

South America'

North America

Ensenadan

Lujanian

Irvingtonian'

Rancholabrean2

N

x ± SE OR

CV N

x ± SE

OR

CV N

X

± SE

OR

CV

N

x ± SE

OR

CV

C AP 2

42.5 - 42.0-43.0

- 13

44.6 ± 1.240

37.6-51.3

10.6

24

42.1 ± 0.537

36.0-46.1

6.25

T 2

19.1 - 18.8-19.3

13

21.5 ± 0.103

20.3-24.0

6.27

24

20.6 ± 0.374

18.0-22.9

8.89

P3 AP -

- - -

- 9

17.2 ± 0.616

13.7-20.5

10.74

17

17.1 ± 0.329

15.0-19.7

7.94

T -

_ _ _

9

9.4 ± 0.321

8.4-1 1.5

10.24

17

9.3 ± 0.150

8.4-10.6

6.66

P4 AP -

- - -

- 9

40.5 ± 1.43

38.4-44.4

10.61 1

37.2

-

22

40.1 ± 0.538

33.4-46.0

6.29

T -

— - —

- 9

15.4 ± 0.258

14.2-16.4

5.03 1

15.6

-

22

16.5 ± 0.329

14.2-19.9

9.36

M1 AP -

_ _ _

- 4

5.7 ± 0.286

5. 0-6. 2

10.05

T -

-

- 4

13.4 ± 0.314

12.7-13.9

4.69

C AP 3

14.2 ± 0.38 13.5-14.8

4.58 9

16.6 ± 1.05

15.6-17.8

24.13 3

14.2

± 0.087

14.1-14.4

1.07

22

14.7 ± 0.231

13.0-16.6

7.40

T 2

10.1 - 10.0-10.1

- 9

1 1.2 ± 0.179

10.0-12.3

4.80 3

9.1

± 0.284

8. 5-9.6

6.24

22

10.6 ± 0.413

9.7-12.2

19.22

P, AP 2

12.0 - 11.4-12.5

- 1

6.0 -

-

- 4

8.7

± 1.02

5.7-10.1

23.40

2

6.2 ± -

5. 7-7.0

—

T 2

7.4 - 7. 3-7. 5

- 1

6.0 -

-

- 4

6.3

± 0.450

5. 0-7.0

14.52

2

8.1 ± -

6. 5-9. 6

-

P4 AP 4

25.5 ± 0.50 24.2-26.5

3.94 10

27.7 ± 0.426

26.3-29.7

4.87 3

26.1

± 1.47

24.0-28.9

9.72

25

24.6 ± 0.273

22.5-26.8

5.55

T 4

10.9 ± 0.256 10.2-11.4

4.70 9

12.3 ± 0.206

1 1.5-13.5

5.08 3

11.1

± 0.550

10.1-12.0

8.59

23

1 1.9 ± 0.173

10.9-12.8

6.97

M, AP 3

26.3 ± 1.63 24.5-29.5

10.73 10

28.3 ± 0.445

25.7-30.0

4.98 4

28.1

± 0.564

27.0-29.5

4.01

25

28.7 ± 0.338

25.9-32.1

5.89

T 3

13.9 ± 0.874 12.2-15.0

10.89 10

13.3 ± 0.137

13.0-14.2

3.26 4

13.1

± 0.147

12.2-15.0

9.88

25

14.3 ± 0.236

12.8-16.1

8.28

2C. Limb length measurements

South America1

North America2

Ensenadan

Lujanian

Rancholabrean

N x ± SE

OR

CV

N

x ± SE

OR

CV

N

it ± SE

OR

CV

Humerus

2 334.0 -

316.0-352.0

_

6 365.4 ± 10.50

340.5-395.0

7.04

10

344.9 ± 7.80 309.0-385.0

7.13

Radius

1 225.0 -

—

-

5 275.7 ± 8.06

249.7-294.8

6.54

10

265.7 ± 6.84 235.0-295.0

8.14

Metacarpal III

1 95.0 -

-

-

3 95.4 ± 1.10

94.3-97.6

2.00 737

-

- 83.0-107.8

-

Femur

4 352.8 ±2.81

347.0-360.0

1.59

4 384.0 ±9.17

365.0-408.0

4.78

iO

367.7 ± 9.80 317.0-408.0

8.43

Tibia

7 251.4 ±7.86

220.0-282.6

8.27

5 280.4 ±4.21

272.5-294.2

3.36

10

273.6 ± 8.10 239.0-305.0

9.31

Metatarsal III

1 98.5 -

_

4 98.0 ± 0.595

94.7-99.1

1.21 766

_

85.5-1 1 1.5

_

Contributions in Science, Number 370

Berta: Smilodon in the Americas 15

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life

Primed at Alien Press. Inc., Lawrence, Kansas

SO

A NEW RECORD OF GIANT SHORT-FACED BEAR,
ARCTODUS SIMUS, FROM WESTERN NORTH AMERICA
WITH A RE-EVALUATION OF ITS PALEOBIOLOGY

Steven D. Emslie1 and Nicholas J. Czaplewski2

ABSTRACT. Remains of the extinct giant short-faced bear, Arctodus
simus, from Labor-of-Love Cave in Nevada constitute the first rec-
ord of this species in eastern Nevada. The skeleton was found in
association with black bear remains, Ursus americanus, eroding from
a gravel lens in silt sediments along the banks of a stream flowing
through the cave. The deposits are probably late Pleistocene in age.
The Arctodus remains are from a small adult and represent one of
the most complete skeletons recovered of this species. Association
of cranial with postcranial bones allowed us to determine proportions
and ratios of limb elements in a single individual. The hypothesis
that A. simus was a highly predaceous cursorial carnivore is exam-
ined. We prefer to believe that this animal was largely herbivorous,
based upon comparative studies of the skull and functional mor-
phology of the jaw in living Tremarctos ornatus, and upon the non-
cursorial features in the limbs of A. simus.

INTRODUCTION

The recovery of a partial skeleton of the giant short-faced
bear, Arctodus simus, and the black bear, Ursus americanus,
from a cave in east-central Nevada, represent the first records
of these bears from eastern Nevada; U. americanus currently
occurs in western Nevada (Hall, 1946). The cave and the
bear remains were originally discovered by members of a
local speleological society in Ely, Nevada, who named the
cave “Labor-of-Love Cave.” Other mammals represented at
this cave include Ursus cf. arctos and Ovis canadensis.

The cave is located at the base of a limestone cliff on the
east side of the Schell Creek Range, White Pine County,
Nevada. The east-facing entrance is at an elevation of 2050
m. The cave extends over 137 m into the mountain along a
fault plane and consists of one main passage. Currently, a
small stream flows the entire length of the cave and emerges
at the base of a talus slope 30 m below the cave entrance.
Air and water temperatures in the cave remain constant year-
round at 10°C.

Postcranial material from Labor-of-Love Cave includes
the first associated limb elements of A. simus on which rel-
ative proportions can be determined. This analysis, together
with earlier studies on the living spectacled bear, Tremarctos

Contributions in Science, Number 371, pp. 1-12
Natural History Museum of Los Angeles County, 1985

ornatus, an animal similar in structure to A. simus, allows a
re-evaluation of the paleobiology of the latter species.

We conclude that A. simus was not necessarily adapted for
a cursorial, predaceous life-style, nor could it have attained
its large size by strict camivory. Based on skull and jaw
architecture, tooth morphology and wear, and limb structure,
A. simus was probably largely herbivorous, but retained bone-
crushing capabilities and may have been an opportunistic
predator and scavenger.

SITE LOCATION

Two concentrated bone deposits (BC-1, BC-2) were found
in two pools of water along the stream edge, and located
approximately 10 m apart at the rear of the cave (Fig. la).
Diversion of stream flow through fissures in the limestone
bedrock maintains these pools as quiet backwater areas.

In summer 1982, under an emergency permit from the
U.S. Forest Service, one of us (SDE) collected and preserved
bones from the two pools before they were washed away by
fluctuations in stream flow or stolen by vandals. The permit
was authorized under the aegis of the Natural History Mu-
seum of Los Angeles County (LACM) where the specimens
are now housed. Bones in the cave were photographed,
mapped, and collected over a period of three days. Spatial
orientation of bones was mapped (Figs, lb, 2) by triangu-
lation, using two or three large aluminum nails, placed in the
substrate at each bone concentration. Each bone was then
carefully removed, wrapped in aluminum foil to avoid con-
tamination, assigned a field number, and packed in a labeled
box prior to removal from the cave. Excavations in cave
strata were not initiated, and are planned for future inves-
tigations. Consequently, no detailed analyses of cave sedi-
ments have been completed. The material was transported

1. Department of Zoology, University of Florida, Gainesville,
Florida 3261 1.

2. Department of Biological Sciences, Northern Arizona Uni-
versity, Flagstaff, Arizona 860 1 1 .

ISSN 0459-8113

enlarged
in Fig, 2

X

a

50 m

Figure 1. Plan view (a) and cross section (b) of Labor-of-Love Cave showing the locations of Bone Concentrations (BC) #1 and #2 and the
gravel lens from which the bones at BC-2 were eroding.

to the George C. Page Museum of LACM for curation and
transferred to LACM for permanent storage. The bones were
identified using comparative collections of Pleistocene and
Recent mammals in the museum. Other fossil remains of A.
simus at the American Museum of Natural History, New
York, and Recent remains of Ursus arctos and U. americanus
at the U.S. National Museum, Washington, D.C., were also
examined.

All vertebrate fossils recovered from the cave are identified
below, but only the remains of A. simus are discussed in
detail in this paper.

SYSTEMATIC PALEONTOLOGY
Order Carnivora
Family Ursidae
Genus Urs us Linnaeus, 1758
Ursus americanus Pallas, 1780

(Black Bear)

REFERRED MATERIAL. LACM 122435, several skull
fragments including maxillae and premaxillae, both lower
jaws, atlas, axis, one thoracic and three other vertebral frag-

ments, both scapulae, humeri, ulnae, radii, a right tibia, meta-
carpal II (right and left), III (right), IV (right and left), and
V (right and left), metatarsal III (right) and IV (right), and
ten rib fragments; LACM 123780, left tibia; LACM 123783,
diaphyses of right and left humeri; LACM 123786, right M,.

DISCUSSION. LACM 122435 was recovered from BC-
1 and represents an immature individual, approximately 3-
4 years old at times of death based on epiphyseal fusion of
long bones. LACM 123780 was also found in BC-1 but rep-
resents another individual. LACM 123783 and 123786 were
found eroding from the gravel lens at BC-2, in direct asso-
ciation with bones of A. simus (Fig. 2b, c). A minimum of
three U. americanus individuals are represented.

Ursus cf. U. arctos Linnaeus, 1758

(Grizzly Bear)

REFERRED MATERIAL. LACM 123782, diaphysis of
left tibia.

DISCUSSION. This tibia, found at BC-1, compares well
in size and morphology to that of a modem Kodiak bear
(Fig. 3). It is more robust, with a shaft flared at both proximal
and distal ends, and has a more sharply ridged shaft than in
other bear species. These characters are lacking in late Pleis-

2 Contributions in Science, Number 371

Emslie and Czaplewski: Paleobiology of Arctodus simus

Table 1. Measurements of the right mandible (LACM 122434) of
Arctodus simus from Labor-of-Love Cave, Nevada.

Length, anterior base of C, to condylar

process

253.0

mm

Depth of mandible at P,

56.5

mm

Depth of mandible at coronoid process

ca. 112-113

mm

Alveolus length, C-M,

154.0

mm

Alveolus length, P,-M,

120.5

mm

Alveolus length, P,-M,

111.0

mm

Alveolus length, P,-M,

98.2

mm

Alveolus length, P4-M,

87.0

mm

Alveolus length, M-M,

74.5

mm

Alveolus length, M,-M,

44.2

mm

C,, anterior-posterior length

28.0

mm

greatest width

19.7

mm

M,, anterior-posterior length

30.2

mm

trigonid length

19.8

mm

trigonid width

12.8

mm

talonid width

17.2

mm

M,, anterior-posterior length

28.0

mm

trigonid width

20.1

mm

talonid width

18.3

mm

M3, anterior-posterior length

18.7

mm

greatest width

14.5

mm

tocene remains of A. simus and U. americanus from Labor-
of-Love Cave and Rancho La Brea, and from modem U.
americanus.

Genus A ret odus Leidy, 1854

Arctodus simus (Cope) 1879

(Giant Short-faced Bear)

REFERRED MATERIAL. LACM 1 2378 1, proximal left
femur; LACM 122434, numerous skull fragments including
premaxillae and maxillae, both lower jaws (see Tables 1, 2;
Figs. 4, 5), humeri, radii, ulnae, femora, and tibiae, atlas,
right metacarpal III, right metatarsal V, fibula fragment, pha-
lanx, and one rib fragment.

DISCUSSION. LACM 123781 was recovered from BC-1
and is from an immature individual as it lacks the proximal
epiphysis. LACM 122434 was recovered from BC-2 and rep-
resents one adult individual. A minimum of two A. simus
individuals are represented.

Order Artiodactyla

REFERRED MATERIAL. LACM 123784, fragmented
humerus.

Table 2. Measurements of teeth (right maxilla) of Arctodus simus
(LACM 122434) from Labor-of-Love Cave, Nevada.

Alveolus length, C'-M-

137.6

mm

Alveolus length, P'-M2

100.1

mm

Alveolus length, P2-M2

93.0

mm

Alveolus length, P3-M2

87.3

mm

Alveolus length, P4-M2

78.7

mm

Alveolus length, M'-M2

56.3

mm

C1, anterior-posterior length

27.9

mm

greatest width

17.9

mm

P4, anterior-posterior length

20.8

mm

greatest width

14.7

mm

M1, anterior-posterior length

24.1

mm

greatest width

23.8

mm

M2, anterior-posterior length

35.0

mm

greatest width

21.7

mm

DISCUSSION. This specimen, recovered from BC-2, is
too fragmented for positive identification beyond order, but
is from a deer-sized animal. Its position in the pool at time
of discovery is shown in Figure 2a.

Family Bovidae

Genus Ovis Linnaeus, 1758

Ovis canadensis Shaw, 1 804

(Bighorn Sheep)

REFERRED MATERIAL. LACM 123785, left M3.

DISCUSSION. Recovered from BC- 1 .

GEOLOGIC HISTORY AND TAPHONOMY

Exact geologic age of the fossils is not known; two bones, an
ulna of U. americanus from BC-1 and a humerus of A. simus
from BC-2, were submitted to Beta Analytic, Coral Gables,
Florida, for radiocarbon dating. Dates on collagen of 4160
± 1 80 years B.P. were obtained for the ulna of U. americanus
and 5320 ± 120 years B.P. for the humerus of A. simus
(Murray Tamers, pers. comm.). The unusually young date
for A. simus is inconsistent with other fossil records of this
species. Error in the dates may be due to low amounts of
carbon obtained from the bone or to contaminants in the
water within the cave. All previous finds of A. simus are
older than 12,650 ± 350 years B.P. (Kurten and Anderson,
1980) and are associated with faunas characteristic of the
Pleistocene (Irvingtonian and Rancholabrean Land Mammal
Ages). Based on the number and diversity of complete spe-
leothems we believe that Labor-of-Love Cave has been sealed
since at least the close of the Pleistocene, or approximately
12,000 years B.P., and that the above Cl 4 ages do not reflect
the true ages of this material.

Contributions in Science, Number 371

Emslie and Czaplewski: Paleobiology of Arctodus simus 3

Figure 2. Plan view of the bones at BC-2. All bones shown are from one skeleton of Arctodus simus except (a) Artiodactyla humerus, (b)
Ursus americanus humerus, and (c) U. americanus right M,.

Labor-of-Love Cave developed within mountains formed
primarily of uplifted Dunderberg Shale and limestone of mid-
dle to late Cambrian age (Hose and Blake, 1976). The cave
follows a fault plane through the limestone. Examination of
speleothems, flowstone, and stratified silt deposits indicates
the cave has undergone alternate periods of filling from per-
haps several entrances, erosion of these sediments, and pro-
nounced changes in cave environment. Hanging flowstone
shelves were observed throughout the cave at heights where
water and/or sediments once existed, between 0.5 and 1.5 m
above current levels. Several stalagmites undercut by erosion
are now tilted, and clay sediments still cling to the walls of
the cave near the roof.

All bones at BC-1 were lying submerged at the bottom of
the pool or partially buried in silt in the bottom. Bones at
BC-2 were lying at the bottom of the pool, and on the surface
of a gravel lens at the pool edge (Fig. 2). Undisturbed deposits
nearby show this gravel lens is overlain by a 20-cm-thick
layer of sandy silt, followed by a 25-30-cm-thick layer of
fine silt (Fig. lb). Bones exposed on the gravel lens included
nearly articulated pairs of radii and ulnae and a partially

buried tibia of A. simus (Fig. 6b), and the molar and a left
humerus of U. americanus. The close association and artic-
ulation of the remains of A. simus suggest the gravel lens is
the site of primary deposition of these bones (Fig. 2).

Fine, densely packed laminar cross-bedded deposits are
still present in the rear of the cave (Fig. 6c) indicating that
deposition and erosion of sediments has been occurring over
a long period of time. We believe that the bear skeletons have
eroded from the deposits relatively recently, based on their
completeness and close association at each bone concentra-
tion. Some bones were found where originally deposited, in
the gravel lens at BC-2. Those bones which were eroded from
these sediments probably were transported only a short dis-
tance (<5 m) before being redeposited as found. Both bear
skeletons had fragmented skulls (possibly caused by the pres-
sure of overlying sediments) and corresponding skeletal ele-
ments preserved at each pool. Possible carnivore tooth marks
on the bone suggest the bear carcasses had been scavenged
prior to burial. The absence of pelves or complete scapulae
may support this conclusion as these elements are usually
the first to be destroyed during carnivore feeding (Hill, 1980;

4 Contributions in Science, Number 371

Emslie and Czaplewski: Paleobiology of Arctodus simus

Binford, 1981). Most of the ribs, vertebrae, and phalanges
from these skeletons were not recovered, presumably having
been washed downstream or redeposited in gravel bars. These
elements were found by Voorhies (1969) to be most suscep-
tible to fluvial transportation in experiments with Recent
mammal carcasses.

The relative completeness of the black bear skeleton from
BC-1 and the short-faced bear skeleton from BC-2 suggests
that these animals died in the cave, perhaps during a period
of winter hibernation. Bones of other animals found in the
cave also display breakage and tooth marks, and may rep-
resent food items transported to the cave by carnivores.

PALEOECOLOGY

Detailed information on the paleoecology of eastern Nevada
during the late Pleistocene is available from recent studies
in Smith Creek Canyon, Snake Range, which is located 75
km northeast of Labor-of-Love Cave. The elevation of the
entrance to this canyon is 1710 m and the area is similar in
topography and environment to the area of Labor-of-Love
Cave. Studies of the paleoecology at Smith Creek Canyon
can, therefore, be useful in reconstructing the habitat avail-
able to A. simus in eastern Nevada during the late Pleisto-
cene.

Analysis of packrat middens from three caves in Smith
Creek Canyon provided Thompson (1979) with specific in-
formation on the vegetation of this area during the late Wis-
consinan and early Holocene. His results indicated that many
plants occurred in the canyon during the Pleistocene that are
found only at higher elevations today. These plants included
bristlecone pine ( Pinus longaeva ), fern-bush (Chamaeba-
tiaria millefolium ), gooseberry currant (Ribes montigenum),
cliff-bush (Jamesia americana), and bush oceanspray ( Ho-
lodiscus microphyllus). Spruce ( Picea ) and fir {Abies) were
absent from the samples. Thompson (1979) postulated that
the period 12-1 3,000 years B.P. was characterized by greater
effective moisture than today with conditions changing to
those of the present by 10,500 years B.P.

Thompson ( 1979), Thompson and Mead (1982), and Mead
et al. ( 1 982) have shown that much of the Great Basin during
the late Pleistocene consisted of pluvial lakes. Above lake
level, valleys were dominated by sagebrush (Artemisia tri-
dentata) cover, similar to the area today. Alpine glaciers
extended down to 2800 m in elevation in the Snake Range,
75 km northeast of Labor-of-Love Cave. Vertebrates re-
covered from the Snake Range, Smith Creek Cave (dated at
> 1 2,000 years B.P.) and packrat middens from Smith Creek
Canyon, include species now restricted to higher elevations
or higher latitudes. These include Ochotona princeps, Sper-
mophilus richardsonii, Thomomys talpoides, Phenacomys
intermedius, and Mustela vison. One undescribed bone iden-
tified as Ursus sp. from Smith Creek Cave is a questionable
identification (Miller, 1979).

In summary, the late Pleistocene habitat of eastern Nevada

Figure 3. The partial left tibia (LACM 1 23782) of Ursus cf. arctos
found at BC-1 compared to a tibia of Recent U. arctos middendorfi.
Note the characteristic flaring of the shaft at the proximal and distal
ends, and the distinct ridges in U. arctos and the fossil specimen.

probably consisted of pluvial lakes and open, sagebrush-cov-
ered flats in the valley bottoms. Mountain ranges surrounding
the valleys bore alpine glaciers above 2800 m with subalpine
to alpine faunal and floral elements in the canyons and valley

Contributions in Science, Number 371

Emslie and Czaplewski: Paleobiology of Arctodus simus 5

Figure 4. The right mandible (LACM 122434) of A rctodus simus from Labor-of-LoveCave showing the premasseteric fossa (pm) characteristic
of the subfamily Tremarctinae. The position of this specimen in the pool at time of discovery is shown in Figure 2.

edges. Open forests of bristlecone pine and other conifers
extended to the valley edge.

DISCUSSION

Arctodus simus remains have been reported from fossil lo-
calities throughout North America, particularly western North
America (Kurten and Anderson, 1980). Other Pleistocene
records not reported by Kurten and Anderson include several
elements of a large A. simus from northern Utah (Nelson
and Madsen, 1983), a fragmentary partial skeleton from Duck
Flat, Washoe County, Nevada (D.R. Tuohy, in litt.), nu-
merous cranial and postcranial elements from Natural Trap
Cave, Big Horn County, Wyoming (L.D. Martin, pers.
comm.), an isolated tooth from Adair County, Oklahoma
(Puckette, 1976), a cranium, five vertebrae, and partial pelvis
from Hot Springs Mammoth Site, Fall River County, South
Dakota (L. Agenbroad, pers. comm.), and a skull, femur, and
ten vertebrae from Cueva Quebrada, Val Verde County, Tex-
as (Lundelius, 1984). Isolated material from Rancho La Brea
has permitted a composite construction of the complete skel-
eton of A. simus. An unpublished record of this species from
American Falls, Idaho, consists also of a relatively complete
skeleton, but the postcranial elements are fragmented and
incomplete (Jon Becker, pers. comm.) and are not useful for
proportional analyses. A nearly complete skeleton from Ful-
ton County, Indiana, is currently under analysis by William
Turnbull and associates (pers. comm.). Consequently, the
material from Labor-of-Love Cave constitutes one of the

most complete skeletons from one individual recovered in
North America. This provides more accurate proportional
data on limbs, with which to evaluate the paleobiology of A.

simus.

PALEOBIOLOGY

Kurten (1967), in his singular monograph on Arctodus bears,
stated that A. simus had cursorial adaptations, and was prob-
ably a highly predaceous carnivore. His conclusion was based
primarily on the long limbs, slender leg elements, and “felid-
like” characters of the skull of A. simus when compared to
other ursids. However, we contend that this species may have
been largely herbivorous based upon characteristics in the
skull, limbs, and body size. In addition, anatomic and func-
tional morphologic comparisons with the living spectacled
bear, Tremarctos ornatus, add support to our argument. Fi-
nally, sympatric records of Pleistocene bears suggest that
Arctodus and Ursus species used different feeding strategies.

SKULL

Kurten (1967) described the skull of A. simus as having sev-
eral felid-like characteristics such as a shortened face, large
rostral breadth, high vaulted calvarium, and similar orien-
tation of the zygomatic arches, and he used these convergent
features to advance his hypothesis that A. simus was an active
carnivore. However, these skull features may be related to
large body size and reduction of premolars in bears, and a

6 Contributions in Science, Number 371

Emslie and Czaplewski: Paleobiology of Arctodus simus

comparison to felids is inappropriate. In addition, the same
features are present in the skull of Tremarctos ornatus, the
only living relative of A ret od us (Davis, 1955). The South
American spectacled bear (T. ornatus ) is primarily herbiv-
orous, feeding on bromeliads, leaves, tree bark and fruits,
berries, and cactus fruits and pulp, and only occasionally eats
carrion; reports of these bears killing prey are uncertain (Pey-
ton, 1980).

One important similarity in the jaw of Tremarctos and
Arctodus is the presence of a premasseteric fossa (Fig. 4). In
a detailed examination of the structure of the dentary and
related muscles in T. ornatus, Davis (1955) concluded this
fossa has one of two functions: it is either a depression
accommodating a small cheek pouch formed in the oral
vestibule (as in the mandrill, Papio maimon), or it reflects
modeling in jaw architecture to allow a larger zygo-
maticomandibularis muscle. This muscle, which is impor-
tant in allowing lateral grinding motions of the jaw and aids
in elevating the mandible, is large in herbivorous animals
and small in carnivorous animals (Davis, 1964). Since the
size of the premasseteric fossa is far too small to be a useful
cheek pouch for the size and food requirements of T. ornatus,
it is probably related more to the jaw mechanics to accom-
modate a large zygomaticomandibularis muscle (Davis, 1955).
Davis (1955:43) also points out that “a functional pouch
occurs elsewhere only in social animals that compete for food
(primates), or in animals that forage for food and retreat to
a safe place to masticate it (rodents). Bears fall into neither
of these categories.” In regards to the fossa itself, Davis (p.
44) states that it does not interfere with the trajectory systems
of the skull and “is situated in tissue that contributes little
or nothing to the internal support of the mandible.” A similar
function for the premasseteric fossa can be projected for A.
simus, because its jaw architecture is very similar to the
closely related T. ornatus. Although morphologic associa-
tions between fossil and living animals should be made with
caution, we believe the close affinities of these two bears
permit reliable comparisons.

The teeth of A. simus also suggest a largely herbivorous
diet. Kurten (1967) suggested that A. simus was predaceous
based primarily on the high-crowned nature of the first lower
molar, and the relatively large size of the canines. He noted
that the protoconid and paraconid in the first molar form a
camassial shear with the paracone and metacone of the upper
fourth premolar. He additionally referred to the trenchant
shape of the first lower molar. Such a camassial shape is
characteristic of many carnivorous fissipeds, but we believe
it is not developed in A. simus. Instead, with moderate wear,
the protoconid and paraconid apices of the first molar are
worn to a relatively flat, blunt loph. While the M, of Arctodus
is high-crowned, it is no more trenchant than that of other
bears and, like U. arctos, it wears flat. Moreover, the only
apparent shear facet is worn into the labial surface of the
tooth in the indentation between the talonid and trigonid
where the paracone of M! occludes, not along the labial sides
of the paraconid and protoconid (Fig. 4). This small shear
facet would be relatively inefficient for slicing flesh. The tri-
gonid of M, is tall and conical, somewhat reminiscent of the

Figure 5. The right maxilla (LACM 122434) of Arctodus simus
from BC-2. Its position in the pool at time of discovery is shown in
Figure 2.

bone-crushing posterior premolars of hyaenas. The talonid
forms an offset, small crushing platform that is confluent with
those of the posterior molars. Perhaps the offset nature of
this tooth functions in crushing plants and bone. The M2 is
extremely broad relative to its length (Table 1) and forms,
with the third molar, a large crushing platform as in omniv-
orous bears. Although cusps of canids and felids can wear
flat, particularly in aged animals, the general trend in most
highly predaceous carnivores is to maintain a sharp, shearing
blade on the camassials. Except in very young specimens, all
of the material of A. simus that we have observed (including
that at the Los Angeles County Museum of Natural History
and the American Museum of Natural History) have flat-
tened cusps. In summary, these characters of the teeth in-
dicate to us that A. simus may have been omnivorous or
primarily herbivorous, but had bone-crushing capabilities.

Tremarctos ornatus, like A. simus, has shorter dentaries
and an overall shorter face when compared to Ursus arctos
(Davis, 1955) and is rarely predaceous, as already noted.
Although the polar bear, an obligate carnivore, has camas-
sials that are no more sectorial than those of other living

Contributions in Science, Number 371

Emslie and Czaplewski: Paleobiology of Arctodus simus 7

8 Contributions in Science, Number 371

Emslie and Czaplewski: Paleobiology of Arctodus simus

bears, they do have tall, pointed cusps in the unworn con-
dition, and the teeth show a trend towards homodonty
(DeMaster and Stirling, 1981), indicating possible conver-
gence with carnivorous pinnipeds. Additionally, the lack of
sectorial teeth in the polar bear may simply reflect its rela-
tively late (mid-Pleistocene) derivation from bunodont U.
arctos (Kurten, 1 964). Moreover, while no one would argue
that felids are not highly predaceous with teeth adapted to a
carnivorous diet, such dental morphology does not neces-
sarily mandate camivory; Felis planiceps of southeast Asia
seems to prefer fruit (Goodwin, 1954). Consequently, Kur-
ten’s suggestion that A. simus was highly predaceous based
on skull structure and teeth cannot be substantiated.

LIMB PROPORTIONS

The major value of the A. simus postcranial material from
Labor-of-Love Cave is that it provides an opportunity to
examine relative limb proportions from a single individual.
Table 3 provides measurements of the limb elements of the
partial skeleton of A. simus from BC-2. Length proportions
of limb elements are compared to similar proportions for
several species of Trernarctos, Arctodus, and Ursus (Table
4). Kurten’s proportions for the fore- and hindlimb elements
were determined using composite material from several fossil
specimens. He found little variation in limb proportions
within A. simus, though considerable size variation within
the taxon was noted. Our calculations (Table 4) indicate that
the limb proportions of A. simus are Lower than those given
by Kurten (1967) but still evince considerable divergence
from other bear taxa; forelimb proportions do not overlap
with Recent bears or fossil Trernarctos, and hindlimb pro-
portions overlap only slightly with U. arctos. This divergence
can be attributed, as Kurten concluded, to the relatively lon-
ger limbs of A. simus compared to other bears.

The strongest evidence for cursorial adaptation in A. simus
offered by Kurten (1967) was its relatively long slender legs.
The humerus and femur are longer than the radius and tibia,
respectively. Hildebrand (1982) discusses skeletal and other
adaptations which increase speed in cursorial animals, in-
cluding an increase in limb length. However, the lengthening
of limbs in cursorial animals usually takes place in the distal
limb segments, not the proximal as in A. simus. Longer prox-
imal limb elements are a feature of digging animals, such as
the badger, and of mediportal and graviportal animals, such
as bears (Davis, 1 964). A. simus, however, lacks other char-
acteristics of a digging life-style such as a long olecranon on
the ulna and long claws. Other non-cursorial characteristics
of bears include plantigrade feet (which is noted for A. simus
by Kurten) and a stride with no or scarcely any unsupported
intervals (Hildebrand. 1982).

Figure 7. Skeleton of A. simus with shaded bones indicating ele-
ments recovered from the adult individual represented at BC-2.

If A. simus was largely herbivorous and not a highly pre-
daceous, cursorial carnivore, another explanation for its long
limbs is needed. An interesting comparison can be made with
the maned wolf, Chrysocyon brachyurus, of South America.
This fox inhabits open, tail-grass prairies (Nowak and Pa-
radise, 1983), has extremely long and slender limbs relative
to body size (as in A. simus), is not especially swift nor does
it take swift prey (Hildebrand, 1954), and runs with a loping
gait (John Eisenberg, pers. comm.). The long limbs may be
an adaptation for increased vision over tall ground cover in
an open habitat (Nowak and Paradise, 1983). A similar in-
terpretation may be applied to A. simus in the Great Basin,
where its former habitat is known, but the habitat for this
species throughout its range in the Pleistocene of North
America is largely unknown. It is equally possible that the
longer limbs of A. simus were used in tearing and pulling
down vegetation, including shrubs and small trees, in order
to feed on leaves, fruits or bark.

BODY SIZE

The body size attained by A. simus may also be an indication
of herbivory. Body size is limited by many factors, including
specialization to specific habitats and diet (Eisenberg, 1981).
In terrestrial mammals, herbivores usually have a larger bio-
mass and greater numerical density than carnivores. This
trend is caused by a more restricted energy base available to
carnivores according to Eisenberg (1981) who also discusses
the average size of recent mammalian genera (using head and
body length) in relation to niche specialization. Table 5 sum-
marizes these data in comparison to A. simus. This table
indicates that Recent terrestrial carnivores are limited to a
maximum size by their diet, and that omnivory has allowed

Figure 6. The gravel and silt deposits at BC-2. (a) The grave! lens containing the remains of A. simus (LACM 122434) extends from the
mound of sediments in the right foreground to the kneeling figure. The stream and pool containing bones are to the right; water is flowing in
the direction of the figure, toward the cave entrance, (b) An articulated radius and ulna, and a partly buried tibia of A. simus eroding from
the gravel lens as shown in Figure 2. (c) Finely laminated and cross-bedded sediments just upstream of BC-2 are all that remain of deposits
that once nearly filled the cave.

Contributions in Science, Number 371

Emslie and Czaplewski: Paleobiology of Arctodus simus 9

Table 3. Measurements (in mm) of limb bones of Arctodus simus (LACM 122434) from Labor-of-Love Cave. Breadth reflects the greatest

lateral-medial breadth, and depth reflects the greatest anterior- posterior depth at each point of measurement.

Element

Total

length

Proximal

breadth

Proximal

depth

Transverse
diameter
prox. head

Least

breadth

shaft

Least

depth

shaft

Distal

breadth

Distal

depth

Left ulna

375

28.8

65.2

—

—

—

—

__

Left radius

355

—

37.9

—

26.2

17.1

—

26.0

Right humerus

454

79.5

-

97.7

35.3

33.9

—

62.8

Right femur

490

124.0

58.0

62.7

39.6

30.4

99.0

56.2

Right tibia

352

90.0

81.7

-

31.2

34.0

63.6

40.5

Right V metatarsal

85

22.5

23.3

—

11.7

10.5

18.3

15.3

Right III metacarpal

—

17.5

31.1

—

13.5

13.1

—

-

bears to attain larger average sizes. The largest living bear is
the Kodiak Bear, U. a. middendorfi, with a mean biomass
of approximately 300 kg (Eisenberg, 1981: Appendix 2), al-
though individual bears may be as large as 700 kg (Novak
and Paradiso, 1983). It should be pointed out that the giant
panda ( Ailuropoda melanoleuca), a herbivore, reaches a bio-
mass of approximately 182 kg and a head and body length
of 1200-1300 mm, indicating an herbivorous diet does not
necessarily result in a large body size.

In comparing measurements of A. simus (as estimated by
Kurten, 1967), to Eisenberg’s data (Table 5) and our own, it
is apparent that this species exceeded the modem limitations
for a terrestrial carnivore. Only two Recent skeletons of U.
arctos (USNM 216206 and USNM 199252, Alaskan males)

equal or exceed the size of A. simus from Labor-of-Love
Cave, which we believe to be a small adult, perhaps a female,
compared to other finds. A recent record of A. simus from
Utah, representing the largest known individual of this species,
was estimated by Nelson and Madsen (1983) to have been
25-30% larger than all previous records, and to have weighed
between 620-660 kg. This estimate is over four times the
average size of the largest terrestrial carnivore ( Panthera )
today. These measurements place A. simus well within the
size range of modem (and Pleistocene) terrestrial frugivore/
omnivores and herbivore/grazers as presented by Eisenberg.
If we accept Kurten’s interpretation of A. simus as having
been primarily carnivorous, then this animal was atypical in
size. It is more logical to assume that A. simus reached its

Table 4. Comparison of relative proportions of limb segments of Arctodus simus, Tremarctos floridanus, T. ornatus, Ursus arctos, and U.
americanus using greatest length for each element, in mm.

Arctodus simus

Radius

Humerus

x 100

N

x

Range

Tibia

Femur

x 100

N

x

Range

Labor-
Hay of-Love

Male1 Female1 Springs1 Cave

T. floridanus'1

Male Female T. ornatus 2

1/2 5/6

87.6 85.6

1/1 3/3 2/3 4/4

78.2 83.0 83.8 84.3

- - - 82.8-86.0

7/7 1/1 1/1

74.5 74.8 71.4

3/3 7/7 4/4

73.4 74.0 79.5

_ _ 74.8-81.7

U. arctos U. americanus

9/9 5/5

88.4 90.3

83.0-95.0 85.5-97.9

9/9 5/5

73.5 77.0

70.4-75.1 74.8-78.4

' Data from Kurten (1967); we were unable to repeat his calculation of the male R/H ratio from his Table 27 and have changed this value
according to data he provides in Tables 1 0 and 1 1 . However, because these ratios are based on non-associated elements from different localities,
their accuracy is questionable.

2 Data from Kurten (1966).

10 Contributions in Science, Number 371

Emslie and Czaplewski: Paleobiology of Arctodus simus

great size by being primarily omnivorous or herbivorous. We
have demonstrated here that the hypothesis for herbivory is
equally plausible to one of camivory, but neither hypothesis
can be rejected with an analysis of functional morphology.
We prefer to believe that A. simus was primarily herbivorous,
as T. ornatus. but also may have been an opportunistic pred-
ator and scavenger with bone-crushing capabilities.

SYMPATHY OF PLEISTOCENE BEARS

The remains at BC-2 indicate a co-occurrence of A. simus
and U. americanus in this area of Nevada. Sympatric records
of these two bears are not unusual and have been documented
at several Pleistocene localities including Rancho La Brea
and McKittrick (Merriam and Stock, 1925; Schultz, 1938).
However, associations of A. simus and U. arctos are rare and
have occurred only at Little Box Elder Cave in Wyoming
and Fairbanks II in Alaska (Kurten and Anderson, 1974,
1980). Kurten and Anderson (1980) speculate that invading
U. arctos in the late Wisconsinan may have played a role in
the extinction of A. simus through competition. However,
this scenario is at odds with Kurten’s earlier suggestions that
A. simus was a highly predaceous carnivore. If we accept
Kurten’s interpretations it seems unlikely that the larger and
more powerful A. simus could be out-competed by the small-
er and less powerful U. arctos. Finally, apparent sympatry
of U. americanus and A. simus during the late Pleistocene is
more likely if the former species was omnivorous, as it is
today, and the latter species was largely herbivorous. In this
situation, the invasion of a third species, U. arctos, which
can be primarily carnivorous in parts of its range today (No-
wak and Paradiso, 1983), may have allowed a sympatric
relationship between these three species. If further investi-
gations in Labor-of-Love Cave provide more positive evi-
dence for an association of A. simus and U. arctos there,
valuable insights on the interspecific relationship of these
species may be gained.

ACKNOWLEDGMENTS

High Desert Grotto members Sam Baker, Dennis Hone, and
Robert Swain discovered and protected Labor-of-Love Cave
for scientific study. Their contributions to this study are greatly
appreciated and cannot be overemphasized. Fieldwork was
completed under a grant from the Cave Research Founda-
tion. Special thanks are extended to this foundation and to
the U.S. Forest Service which provided funds for C14 anal-
ysis. Fieldwork was accomplished with the assistance of the
High Desert Grotto, Ely, Nevada, Frank Johnson of the For-
est Service, local personnel from the Bureau of Land Man-
agement and Forest Service, and George Jefferson and Les
Marcus from the George C. Page Museum of LACM. The
Page Museum also kindly provided research space and the
use of its facilities and personnel in preparing the material
for study. George Jefferson, Bill Akersten, Chris Shaw, and
Shelly Cox were especially helpful at the museum and pro-
vided many useful suggestions. Phil Angle provided valuable
assistance at the U.S. National Museum, Washington, D.C.
We benefitted greatly from the comments of G.E. Goslow,

Table 5. Comparisons of the macroniche, average length, and weight
of Recent mammalian genera (after Eisenberg, 1981) to the proposed
niche, length (estimated by Kurten, 1967) and weight (estimated by
Kurten, 1967 and Nelson and Madsen, 1983) of Arctodus simus.

Taxon

Niche

'Average
head and
body length
(mm)

2Range of
average
weights
(kg)

Equus

Terrestrial herbivore/

1508

166-260

grazer

A. simus

Terrestrial herbivore/

1432

270-660

omnivore

Ursus

Terrestrial frugivore/

817

77-300

omnivore

Panthera

Terrestrial carnivore

718

39-151

1 Average head and body length for Recent mammalian genera was
derived by Eisenberg (1981, Table 43, p. 265) by summing the mean
values for the two extreme size-classes in a genus.

2 Range of weights for Recent mammalian genera was taken from
weights of all species within a genus as presented by Eisenberg (1981,
Appendix 2).

Jim I. Mead, David Elliott, Chris Shaw, G. Jefferson and

several anonymous reviewers who reviewed an earlier draft

of this paper, and from discussions with John Eisenberg.

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Emslie and Czaplewski: Paleobiology of Arctodus simus 1 1

Hose, R.K., and M.C. Blake, Jr. 1976. Geology and mineral
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Kurten, B., and E. Anderson. 1974. Association of Ursus
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Pittsburgh.

Mead, J.I., R.S. Thompson, and T.R. Van Devender. 1982.
Late Wisconsinan and Holocene fauna from Smith Creek
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Geology, Special Paper 1:1-69.

Accepted 8 July 1985.

12 Contributions in Science, Number 371

Emslie and Czaplewski: Paleobiology of Arctodus simus

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